KR102603683B1 - Plant growth regulatory genes encoding seeds plant specific extensin motif containing proteins and uses thereof - Google Patents
Plant growth regulatory genes encoding seeds plant specific extensin motif containing proteins and uses thereof Download PDFInfo
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Abstract
본 발명은 종자 식물 특이 익스텐신 모티브를 포함하는 단백질을 코딩하는 식물 생장조절 유전자 및 이의 용도에 관한 것으로, 본 발명에 따라 조직/발달단계 비특이적 프로모터와 종자 식물 특이 익스텐신 모티브를 포함하는 단백질 (seeds plant specific extensin motif containing proteins, SEP) 코딩 서열이 연결된 재조합 벡터를 사용하여 지질분해, 글리옥실산 회로, 글리옥시좀/퍼옥시좀 발생 조절이 가능하므로, 식물의 유지 함량과 조성을 제어를 통한 유지식물 및 유지작물의 개량 또는 개발에 유용하게 활용될 수 있다.The present invention relates to a plant growth regulation gene encoding a protein containing a seed plant-specific extensin motif and its use. According to the present invention, a protein containing a tissue/developmental stage non-specific promoter and a seed plant-specific extensin motif is provided. (seeds plant specific extensin motif containing proteins, SEP) It is possible to control lipolysis, glyoxylic acid cycle, and glyoxisome/peroxisome development by using a recombinant vector with linked coding sequence, so it is possible to control the oil content and composition of the plant. It can be useful in the improvement or development of oil plants and oil crops.
Description
본 발명은 종자 식물 특이 익스텐신 모티브를 포함하는 단백질을 코딩하는 식물 생장조절 유전자 및 이의 용도에 관한 것으로, 구체적으로 애기장대 유래 종자 식물 특이 익스텐신 모티브를 포함하는 단백질 (seeds plant specific extensin motif containing proteins, SEP)을 코딩하는 sep 유전자 (atr7, At5G21280) 또는 밀 유래 SEP 단백질을 코딩하는 sep 유전자 (TraesCS5B02G518900.1)를 포함하는 재조합 벡터로 식물세포를 형질전환시켜 sep 유전자의 발현을 조절하는 단계를 포함하는 식물체의 지방산 함량과 조성을 조절하는 방법, 식물체의 생장과 개화시기를 조절하는 방법, 종자발아를 조절하는 방법, 식물세포 크기를 조절하는 방법, 식물체 내 지질분해를 조절하는 방법, 식물체 내 글리옥실산 회로를 조절하는 방법, 글리옥시좀/퍼옥시좀 생산을 조절하는 방법 및 상기 제조된 형질전환 식물체에 관한 것이다.The present invention relates to a plant growth regulation gene encoding a protein containing a seed plant-specific extensin motif and its use, and specifically to a protein containing a seed plant-specific extensin motif derived from Arabidopsis (seeds plant specific extensin motif). A step of controlling the expression of the sep gene by transforming plant cells with a recombinant vector containing the sep gene (atr7, At5G21280) coding for containing proteins (SEP) or the sep gene (TraesCS5B02G518900.1) coding for the wheat-derived SEP protein. A method of controlling the fatty acid content and composition of a plant, including a method of controlling the growth and flowering time of a plant, a method of controlling seed germination, a method of controlling plant cell size, a method of controlling lipid decomposition in a plant, a method of controlling lipid decomposition in a plant, It relates to a method for controlling the glyoxylic acid cycle, a method for controlling glyoxysome/peroxisome production, and the prepared transgenic plant.
익스텐신 (extensin, EXT) 모티브는 세린 1개와 연이어 3~5개의 프롤린으로 구성된 펩티드 (SerPro3~5)로서, 애기장대의 경우 177개의 유전자가 알려져 있다. 이중 42개는 SerPro3~5 모티브 주위에 Tyr-Val-Tyr 모티브를 갖는 익스텐신이다. 11개는 익스텐신 모티브 외에 Leu과 Cys이 반복되는 도메인 (LRR)을 포함하는데 이를 LRR-EXT라 한다. 이외에 아라비노갈락탄 (arabinogalactan) 단백질-EXT (4개), PERK (15개), 포르민 (액신-미세소관 결합도메인, 11개)가 있다. 이들은 세포신장, 신호감지 및 전달, 세포골격 형태조절에 관여하는 것으로 알려져 있다 (Borassi, Cecilia & Sede, Ana & Mecchia, Martin & Salter, Juan & Marzol, Eliana & Muschietti, Jorge & Estevez, Jose. (2015). An update on cell surface proteins containing extensin-motifs. J. Exp. Bot. 67).The extensin (EXT) motif is a peptide (SerPro3~5) composed of one serine followed by three to five prolines. In Arabidopsis, 177 genes are known. Of these, 42 are extensins with Tyr-Val-Tyr motifs around the SerPro3 to 5 motifs. In addition to the extensin motif, 11 contain a Leu and Cys repeating domain (LRR), which is called LRR-EXT. In addition, there are arabinogalactan proteins - EXT (4), PERK (15), and formin (axin-microtubule binding domain, 11). They are known to be involved in cell elongation, signal sensing and transmission, and regulation of cytoskeletal morphology (Borassi, Cecilia & Sede, Ana & Mecchia, Martin & Salter, Juan & Marzol, Eliana & Muschietti, Jorge & Estevez, Jose. (2015 ). An update on cell surface proteins containing extensin-motifs. J. Exp. Bot. 67).
애기장대 유래의 At5g21280(atr7) 유전자는 안토시아닌 (anthocyanin) 생합성 조절에 관여하는 전사인자 PAP1과 프로그램화세포사멸을 유도하는 AAL-toxin 저항성 유전자를 발굴하는 과정에서 보고되었다. 안토시아닌은 식물의 영양 및 생식기관의 색깔, 곤충 등의 꽃가루 매개자 유인, 종자 분산 및 활성산소 등에 의한 손상을 막는 항산화제로 작용한다. R2R3 MYB 전사인자인 PAP1 (production of anthocyanin pigment 1)은 WD-40 단백질인 TTG1 (transparent testa glabra 1) 그리고 bHLH (basic helix-loop-helix) 전사인자인 TT8, GL3 및 EGL3와 함께 MBW (MYB-bHLH-WD40) 전사복합체가 구성되면 안토시아닌 생합성 유전자 발현이 활성화된다. 반면, R3 MYB 전사인자인 MYBL2는 LBW (MYBL2-bHLH-TTG1) 복합체를 형성하여 안토시아닌 생합성을 억제한다. 상기 복합체 이외에 MYB11, 12, 111, 113 및 114는 복합체를 형성하지 않고 독립적으로 안토시아닌 생합성 유전자의 발현을 조절하는 것으로 알려졌다. 안토시안이 과량축적되는 표현형인 PAP1D 모본에서 T-DNA가 At5G21280의 엑손에 삽입될 시 돌연변이 (보라색) 표현형을 나타내어 p296으로 명명되었으나 At5G21280가 안토시아닌의 조절 혹은 식물의 생장과 분화에 미치는 영향은 알려진 바 없다.The At5g21280 (atr7) gene from Arabidopsis was reported in the process of discovering the transcription factor PAP1, which is involved in the regulation of anthocyanin biosynthesis, and the AAL-toxin resistance gene that induces programmed cell death. Anthocyanins act as antioxidants to prevent damage caused by the color of plants' nutritional and reproductive organs, attraction of pollinators such as insects, seed dispersal, and free radicals. PAP1 (production of anthocyanin pigment 1), an R2R3 MYB transcription factor, produces MBW (MYB- When the bHLH-WD40) transcription complex is formed, anthocyanin biosynthetic gene expression is activated. On the other hand, MYBL2, an R3 MYB transcription factor, inhibits anthocyanin biosynthesis by forming the LBW (MYBL2-bHLH-TTG1) complex. In addition to the above complex, MYB11, 12, 111, 113, and 114 are known to independently regulate the expression of anthocyanin biosynthesis genes without forming a complex. When T-DNA is inserted into the exon of At5G21280 in the PAP1D model, which is a phenotype in which anthocyanins are excessively accumulated, a mutant (purple) phenotype is displayed and it is named p296 . However, the effect of At5G21280 on the regulation of anthocyanins or the growth and differentiation of plants is not known. .
At5g21280 돌연변이체는 정상적인 생장조건에서는 잎의 크기나 유식물 생체량 등 바이오매스가 감소하는 표현형을 보이고 다른 생장과 발달에 표현형을 보이지 않지만 프로그램화 세포사멸을 유도하는 AAL-toxin, 캐탈라제 억제제인 아미노트리아졸 (aminotriazole)과 제초제의 일종인 파라콰트(paraquat)에 의해 유도되는 산화스트레스에 내성을 보이는 것으로 알려져 있으며, 이러한 산화스트레스 저항성은 비생물학적 스트레스 관련 전사인자 DREB19, HSFA2, ZAT10 등의 축적과 밀접한 연관이 있는 것으로 알려져 있다. 상기 보고는 At5g21280가 식물의 생장과 항산화 반응에 관여함을 시사하나, 단백질 서열상의 정보로는 그 정확한 작용 메카니즘을 추정하기 쉽지 않을 뿐만 아니라 생장과 발달 그리고 환경변화에 대한 반응과정에서 생리학적 기능을 유추하기 어렵다.The At5g21280 mutant shows a phenotype of decreased biomass, such as leaf size and seedling biomass, under normal growth conditions, and does not show any other growth and development phenotypes, but AAL-toxin, which induces programmed cell death, and amino acid, a catalase inhibitor. It is known to exhibit resistance to oxidative stress induced by aminotriazole and paraquat, a type of herbicide, and this oxidative stress resistance is closely related to the accumulation of abiotic stress-related transcription factors such as DREB19, HSFA2, and ZAT10. It is known that there is. The above report suggests that At5g21280 is involved in plant growth and antioxidant response, but it is not easy to estimate its exact mechanism of action based on information on the protein sequence, and it also exerts physiological functions in the process of growth, development, and response to environmental changes. It's difficult to infer.
그러나, 아직까지 상기 애기장대 유래의 At5g21280을 이용하여 형질전환된 식물체 제조방법 및 그에 따른 식물체에 관해서는 알려진 바가 없다. However, there is still nothing known about the method of producing transformed plants using At5g21280 derived from Arabidopsis thaliana and the resulting plants.
이러한 배경 하에, 본 발명자들은 애기장대 밀에서 종자식물 특이 익스펜신형 단백질 (seeds plant specific extensin motif containing proteins, SEP)을 코딩하는 sep 유전자를 새롭게 발굴하였으며, 상기 sep 유전자에 변이가 있는 경우 식물체가 잘 자라지 못하여 생장 및 개화시기가 지연되는 것을 확인하고, 상기 sep 유전자에 변이가 있는 식물체에 sep 유전자를 과발현시키는 경우, 정상적으로 생장 및 개화하는 것을 확인함으로써, 본 발명을 완성하였다.Under this background, the present inventors newly discovered a sep gene encoding seeds plant specific extensin motif containing proteins (SEP) in Arabidopsis wheat, and found that when there is a mutation in the sep gene, the plant grows well. The present invention was completed by confirming that growth and flowering time were delayed due to failure to grow, and that when the sep gene was overexpressed in a plant with a mutation in the sep gene, it grew and flowered normally.
본 발명의 하나의 목적은 sep 유전자를 포함하는 식물체 생장 조절용 조성물을 제공하는 것이다.One object of the present invention is to provide a composition for controlling plant growth containing the sep gene.
본 발명의 다른 목적은 상기 sep 유전자를 포함하는 재조합 벡터를 제공하는 것이다.Another object of the present invention is to provide a recombinant vector containing the sep gene.
본 발명의 다른 목적은 식물체 내에서 sep 유전자를 과발현시키는 단계를 포함하는, 식물체의 생장 촉진 방법을 제공하는 것이다.Another object of the present invention is to provide a method for promoting plant growth, which includes the step of overexpressing the sep gene in the plant.
본 발명의 다른 목적은 식물체 내에서 sep 유전자의 발현을 저해시키는 단계를 포함하는, 식물체의 생장 저해 방법을 제공하는 것이다.Another object of the present invention is to provide a method for inhibiting plant growth, comprising the step of inhibiting the expression of the sep gene in the plant.
이하에서는, 본 발명을 더욱 상세히 설명한다. 한편, 본 발명에서 개시되는 각각의 설명 및 실시형태는 각각의 다른 설명 및 실시 형태에도 적용될 수 있다. 즉, 본 발명에서 개시된 다양한 요소들의 모든 조합이 본 발명의 범주에 속한다. 또한, 하기 기술되는 구체적인 서술에 의하여 본 발명의 범주가 제한된다고 할 수 없다. 또한, 당해 기술분야의 통상의 지식을 가진 자는 통상의 실험만을 사용하여 본 발명에 기재된 본 발명의 특정 양태에 대한 다수의 등가물을 인지하거나 확인할 수 있다. 또한, 이러한 등가물은 본 발명에 포함되는 것으로 의도된다. Below, the present invention is described in more detail. Meanwhile, each description and embodiment disclosed in the present invention may also be applied to each other description and embodiment. That is, all combinations of the various elements disclosed in the present invention fall within the scope of the present invention. Additionally, it cannot be said that the scope of the present invention is limited by the specific description described below. Additionally, those skilled in the art will recognize, or be able to ascertain using no more than routine experimentation, many equivalents to the specific embodiments of the invention described herein. Additionally, such equivalents are intended to be encompassed by this invention.
상기 과제를 해결하기 위한 본 발명의 하나의 양태는 sep 유전자를 포함하는 식물체 생장 조절용 조성물을 제공한다.One aspect of the present invention to solve the above problem provides a composition for controlling plant growth containing the sep gene.
본 발명에서, 용어 "종자식물 특이 익스펜신형 단백질 (seeds plant specific extensin motif containing proteins, SEP)"은 쌍자엽과 단자엽등 종자식물의 핵 단백질로서 비결정형 구조 (intrinsically disordered proteins) 일종이다. 생화학적 기능은 알려지지 않았으며, 돌연변이체 표현형 분석을 통해서 항산화 메커니즘에 관여하는 것으로 알려졌다. 본 발명에 있어서, 상기 sep 유전자는 SEP 단백질을 코딩하는 유전자일 수 있다.In the present invention, the term "seeds plant specific extensin motif containing proteins (SEP)" is a type of intrinsically disordered protein that is a nuclear protein of seed plants such as dicots and monocots. Its biochemical function is unknown, but it is known to be involved in antioxidant mechanisms through mutant phenotype analysis. In the present invention, the sep gene may be a gene encoding a SEP protein.
상기 sep 유전자는 애기장대 또는 밀 유래일 수 있고, 구체적으로 상기 sep 유전자는 서열번호 1의 염기서열을 포함하거나 이로 표시되는 애기장대 유래 At5G21280 유전자위 (atr7 유전자) 또는 서열번호 2의 염기서열을 포함하거나 이로 표시되는 밀 유래 TraesCS5B02G518900.1일 수 있다. 또한 상기 유전자 서열의 상동체가 본 발명의 범위 내에 포함될 수 있다. 상동체의 염기서열은 변화될 수 있으나, 서열번호 1 또는 서열번호 2와 유사한 기능적 특성을 가질 수 있다. 구체적으로, 상기 sep 유전자의 염기서열은 서열번호 1 또는 서열번호 2의 염기서열과 각각 70% 이상, 더 바람직하게는 80% 이상, 더욱 바람직하게는 90% 이상, 가장 바람직하게는 95% 이상, 100% 미만의 서열 상동성을 가지는 염기서열을 포함할 수 있다. 폴리뉴클레오티드에 대한 "서열 상동성의 %"는 두 개의 최적으로 배열된 서열과 비교 영역을 비교함으로써 확인되며, 비교 영역에서의 폴리뉴클레오티드 서열의 일부는 두 서열의 최적 배열에 대한 참고 서열(추가 또는 삭제를 포함하지 않음)에 비해 추가 또는 삭제(즉, 갭)를 포함할 수 있다.The sep gene may be derived from Arabidopsis thaliana or wheat, and specifically, the sep gene includes the base sequence of SEQ ID NO: 1 or the At5G21280 locus (atr7 gene) derived from Arabidopsis represented by this or the base sequence of SEQ ID NO: 2. Or it may be wheat-derived TraesCS5B02G518900.1, which is represented by this. Additionally, homologs of the above gene sequence may be included within the scope of the present invention. The base sequence of the homolog may change, but may have functional properties similar to SEQ ID NO: 1 or SEQ ID NO: 2. Specifically, the base sequence of the sep gene is at least 70%, more preferably at least 80%, more preferably at least 90%, and most preferably at least 95% of the base sequence of SEQ ID NO: 1 or SEQ ID NO: 2, respectively. It may include base sequences with less than 100% sequence homology. The “% sequence homology” for a polynucleotide is determined by comparing a comparison region with two optimally aligned sequences, where a portion of the polynucleotide sequence in the comparison region is a reference sequence (additions or deletions) for the optimal alignment of the two sequences. may contain additions or deletions (i.e. gaps) compared to those that do not contain .
상기 sep 유전자는 서열번호 3 또는 서열번호 4의 아미노산 서열을 포함하는 SEP 단백질을 코딩할 수 있다.The sep gene may encode a SEP protein containing the amino acid sequence of SEQ ID NO: 3 or SEQ ID NO: 4.
상기 "식물체 생장 조절"은 다음 중 선택되는 어느 하나 이상의 특징을 가지는 것일 수 있으나, 이에 제한되지 않는다.The “plant growth control” may have one or more characteristics selected from the following, but is not limited thereto.
i) 식물체의 지방산 함량 조절;i) Control of fatty acid content in plants;
ii) 식물체 내 글리옥시좀 또는 퍼옥시좀 발현 조절;ii) regulation of glyoxisome or peroxisome expression in plants;
iii) 식물세포의 크기 조절;iii) regulation of plant cell size;
iv) 식물체 내 지질 분해 조절;iv) regulation of lipid degradation in plants;
v) 식물체 내 글리옥실산 회로 관련 유전자 발현 조절;v) Regulation of glyoxylic acid cycle-related gene expression in plants;
vi) 식물체의 개화시기 조절; 및vi) Control of flowering time of plants; and
vii) 식물체의 산화스트레스 저항성 조절.vii) Control of oxidative stress resistance in plants.
상기 i) 식물체의 지방산 함량 조절과 관련하여, 본 발명의 일 구현예에서, 애기장대 야생종 (Col-0), At5F21280 유전자에 대한 T-DNA 삽입 돌연변이체 (Salk_006796) 및 At5F21280 유전자를 포함하는 벡터로 형질전환되어 sep 유전자가 보상발현된 Salk_006796 (AtCO)를 암조건에서 7일 생육하고 총 지방산 함량을 확인한 결과, Salk_006796 (sep)의 총 지방산 함량은 Col-0에 비해 약 377% 높았으며, AtCO의 총 지방산 함량은 Col-0와 유사하였다 (도 4b). 이는 sep 유전자 발현을 조절함으로써 식물체 내 지방산 함량을 조절할 수 있음을 시사한다.With regard to i) controlling the fatty acid content of the plant, in one embodiment of the present invention, a wild type of Arabidopsis thaliana (Col-0), a T-DNA insertion mutant for the At5F21280 gene (Salk_006796), and a vector containing the At5F21280 gene are used. Salk_006796 (AtCO), which was transformed and had compensatory expression of the sep gene, was grown under dark conditions for 7 days and the total fatty acid content was checked. As a result, the total fatty acid content of Salk_006796 ( sep ) was about 377% higher than that of Col-0, and that of AtCO Total fatty acid content was similar to Col-0 (Figure 4b). This suggests that fatty acid content in plants can be controlled by regulating sep gene expression.
본 발명에서, 상기 "지방산"은 식물체 내 포함되어 있는 지방산이라면 제한되지 않으나, 예를 들면, C16, C18, 및 C20로 이루어지는 군으로부터 선택되는 지방산일 수 있다.In the present invention, the “fatty acid” is not limited as long as it is a fatty acid contained in the plant, but may be, for example, a fatty acid selected from the group consisting of C16, C18, and C20.
상기 ii) 식물체 내 글리옥시좀 또는 퍼옥시좀 발현 조절 및 iv) 식물체 내 지질 분해 조절과 관련하여, 본 발명의 일 구현예에서, Salk_006796 (sep)은 포도당 처리한 배지에서는 Col-0 대비 각각 90% 수준으로 회복된 반면, 과당, 이당류, 오스모티쿰인 소비톨과 만니톨 처리한 배지에서는 생장 회복현상이 나타나지 않았다 (도 4c). 이는 종자에 저장된 지질은 글리옥시좀/퍼옥시좀이 관여하는 분해대사 경로를 통해서 하배축을 포함하는 생장축 (growing axis)에서 필요로 하는 탄소원과 에너지원으로 사용됨에 따라, Salk_006796 (sep) 돌연변이 식물에서 유적의 축적에 의해서 포도당 신생성이 억제되고, 결과적으로 하배축 등 유식물이 억제되었기 때문에 외부에서 포도당 공급 시 돌연변이체 생장이 야생종 수준으로 회복된 것으로, SEP가 저장 지질 분해에 관여하는 것을 나타내고, 나아가 sep 발현을 조절함으로써 지방산 분해 의존성 에너지 및 탄소대사를 통해서 세포길이 조절이 가능함을 시사한다. In relation to the above ii) regulation of glyoxisome or peroxisome expression in plants and iv) regulation of lipid degradation in plants, in one embodiment of the present invention, Salk_006796 ( sep ) has 90% compared to Col-0 in glucose-treated medium. % level, while growth recovery did not occur in media treated with fructose, disaccharide, Osmoticum sorbitol, and mannitol (Figure 4c). This is because the lipids stored in the seeds are used as carbon and energy sources required by the growing axis, including the hypocotyl, through a metabolic metabolic pathway involving glyoxysomes/peroxisomes, Salk_006796 ( sep ) mutant plants. Since gluconeogenesis was suppressed by the accumulation of lipids, and as a result, seedlings such as hypocotyls were suppressed, the growth of the mutant was restored to the level of the wild species when glucose was supplied from the outside, indicating that SEP is involved in the breakdown of storage lipids. Furthermore, it suggests that cell length can be controlled through fatty acid breakdown-dependent energy and carbon metabolism by regulating sep expression.
본 발명에서, 용어 "글리옥시좀 (glyoxysome)"은 많은 고등식물의 오일을 많이 갖고 있는 종자 내에서 단일막으로 둘러싸여 지방산의 베타 산화 (β-oxidation)와 글리옥실산 (glyoxylate)의 대사를 위해 전문화된 미소체 (microbody)이다. 퍼옥시좀 (peroxisome)에서 특화된 소기관이다.In the present invention, the term "glyoxysome" refers to the term "glyoxysome", which is surrounded by a single membrane within the oil-rich seeds of many higher plants for beta-oxidation of fatty acids and metabolism of glyoxylate. It is a specialized microbody. It is a specialized organelle in the peroxisome.
본 발명에서, 용어 "퍼옥시좀 (Peroxisome)"은 진핵세포의 세포질 내에 존재하는 세포 소기관으로, 리소좀 (lysosome)과 유사하게 막에 둘러싸인 구조물이다. 식물 세포 종자발아 및 영양생장기에 엽록체와 미토콘드리아와 협동적으로 지질분해, 옥신, 지베렐린, 자스몬산을 포함하는 식물호르몬 생합성, 및 광호흡 경로에 관여하며, 세포내 소포체 (ER)에서 출아 (budding)된 후 전구체 프리퍼옥시좀 (preperoxisome)의 확장과 분열에 의해서 성숙되는데 이는 페록신 (peroxin, PEX) 단백질에 의해서 매개된다. 퍼옥시좀의 수, 구조 및 크기는 일정하지 않고 발달단계나 환경변화에 따라서 가변적이며, 펙소파지 (pexophagy)에 의해서 선택적으로 분해된다.In the present invention, the term “peroxisome” is an organelle present in the cytoplasm of eukaryotic cells, and is a membrane-enclosed structure similar to a lysosome. During plant cell seed germination and vegetative growth, it cooperates with chloroplasts and mitochondria to participate in lipolysis, plant hormone biosynthesis including auxin, gibberellin, and jasmonic acid, and the photorespiration pathway, and is budding in the intracellular endoplasmic reticulum (ER). It matures by expansion and division of the precursor preperoxisome, which is mediated by the peroxin (PEX) protein. The number, structure, and size of peroxisomes are not constant and vary depending on the developmental stage or environmental changes, and are selectively decomposed by pexophagy.
상기 iii) 식물세포의 크기 조절과 관련하여, 본 발명의 일 구현예에서, Col-0, Salk_006796 (sep) 및 AtCO의 유식물 하배축 세포크기를 측정한 결과, Salk_006796 (sep)는 Col-0에 비해서 하배축 표피 장축 길이가 54% 수준으로 감소하였으나 AtCO에서는 77-80% 수준의 길이를 보였다 (도 4a 아래). 이는 sep 유전자 발현을 조절함으로써 식물세포의 크기를 조절할 수 있음을 시사한다.With regard to the above iii) control of the size of plant cells, in one embodiment of the present invention, as a result of measuring the cell size of the seedling hypocotyl of Col-0, Salk_006796 ( sep ) and AtCO, Salk_006796 ( sep ) is in Col-0. In comparison, the hypocotyl epidermal long axis length was reduced to 54%, but AtCO showed a length of 77-80% (bottom of Figure 4a). This suggests that the size of plant cells can be controlled by regulating sep gene expression.
상기 v) 식물체 내 글리옥실산 회로 관련 유전자 발현 조절과 관련하여, 본 발명의 일 구현예에서, Salk_006796 (sep)에서 TAG 모빌라이제이션 (SDP1), 베타 산화 (PXA1, ACX2-4, KAT2), 글리옥실산 회로 (CSY3, ACO3, MLS, PMDH1) 그리고 퍼옥시좀 생성 (PEX11A, D, E)과 관련된 유전자량이 야생종에 비해서 2~8배 정도 감소하였으며, 이러한 유전자량 감소는 AtCO에서는 Col-0 수준으로 회복되었다 (도 5). 이는 sep 유전자 발현을 조절함으로써 글리옥실산 회로 관련 유전자의 발현을 조절할 수 있음을 시사한다.Regarding v) regulation of gene expression related to the glyoxylic acid cycle in plants, in one embodiment of the present invention, in Salk_006796 ( sep ), TAG mobilization (SDP1), beta oxidation (PXA1, ACX2-4, KAT2), The amount of genes related to the glyoxylate cycle (CSY3, ACO3, MLS, PMDH1) and peroxisome production (PEX11A, D, E) was reduced by 2 to 8 times compared to the wild species, and this decrease in gene amount was caused by Col-0 in AtCO. level was restored (Figure 5). This suggests that the expression of glyoxylate cycle-related genes can be controlled by regulating sep gene expression.
본 발명에서, 용어 "글리옥실산 회로 (glyoxylate cycle)"는 시트르산 회로 (TCA 회로)의 변형 회로로서 식물과 세균, 원생생물, 균류 등에 존재하는 동화 (anabolic) 경로이다. 글리옥실산 회로에서 시트르산 회로와 차이가 나는 반응은 이소시트르산으로부터 숙신산과 글리옥실산이 생성되는 것과 아세틸-CoA 일부가 글리옥실산과 반응하여 말산이 생성되는 것이다. 글리옥실산 회로는 시트르산 회로의 탈탄산 (decarboxylation) 반응을 우회하는 경로로서, 식물과 미생물이 아세트산으로부터 탄수화물을 합성하고 에너지를 생산할 수 있도록 하는 경로이다. In the present invention, the term “glyoxylate cycle” is a modified cycle of the citric acid cycle (TCA cycle) and is an anabolic pathway that exists in plants, bacteria, protists, fungi, etc. The reaction in the glyoxylic acid cycle that differs from the citric acid cycle is that succinic acid and glyoxylic acid are produced from isocitric acid, and a portion of acetyl-CoA reacts with glyoxylic acid to produce malic acid. The glyoxylic acid cycle is a pathway that bypasses the decarboxylation reaction of the citric acid cycle and allows plants and microorganisms to synthesize carbohydrates from acetic acid and produce energy.
식물에서 글리옥실산 회로가 진행되는 장소는 글리옥시솜 (glyoxysome)이라 불리우는 특별한 미소체 구획이다. 가장 대표적인 글리옥시좀 연관 발생 과정으로 종자의 발아 과정을 들 수 있다. 종자는 주로 트리글리세리드 (triglyceride; 중성지방의 기본 단위) 형태로 지방을 저장하고 있다가, 종자가 발아를 시작하게 되면, 저장 지방로부터 발아 과정에서 필요한 에너지와 탄소원을 공급하기 위한 일련의 대사과정을 진행하게 되는데, 이 과정의 중심에 글리옥실산 회로가 존재한다. 광합성 등을 통해 에너지를 공급받지 못하는 종자가 발아해서 유식물로 생장하고 발달하기 위한 과정에서 필요한 에너지를 공급하기 위하여, 트리글리세리드가 분해되고, 글리옥실산 회로 및 포도당신생합성 (gluconegenesis) 과정을 거치게 되면 식물이 직접 이용가능한 에너지원인 포도당, 설탕 등의 탄수화물이 생성된다.In plants, the site of the glyoxylate cycle is a special microsomal compartment called a glyoxysome. The most representative glyoxysome-related developmental process is the seed germination process. Seeds mainly store fat in the form of triglyceride (the basic unit of neutral fat), and when the seed begins to germinate, it undergoes a series of metabolic processes to supply energy and carbon sources needed during the germination process from the stored fat. At the center of this process is the glyoxylic acid cycle. In order to supply the energy needed for seeds that do not receive energy through photosynthesis to germinate and grow and develop into seedlings, triglycerides are decomposed and go through the glyoxylic acid cycle and gluconeogenesis. Carbohydrates such as glucose and sugar, which are energy sources that plants can directly use, are produced.
상기 vi) 식물체의 개화시기 조절과 관련하여, 본 발명의 일 구현예에서, Salk_006796 (sep)는 37일 생육한 식물체에서 꽃대 및 꽃의 발달이 야생형에 비해 지연되었으나, 이러한 발아 및 생장 지연 표현형은 AtCO에서 sep 도입에 의해 야생형 수준으로 회복되었다 (도 3c). 상기 결과는 생식 단계에서 sep 유전자에 의해 식물체의 종자 발아 지연과 왜소성 및 개화 조절이 가능함을 시사한다. Regarding vi) control of the flowering time of the plant, in one embodiment of the present invention, Salk_006796 ( sep ) showed delayed development of peduncles and flowers in plants grown for 37 days compared to the wild type, but this delayed germination and growth phenotype It was restored to wild-type levels by introduction of sep in AtCO (Figure 3c). The above results suggest that delay in seed germination, dwarfism, and flowering control of plants is possible by the sep gene in the reproductive stage.
상기 vii) 식물체의 산화스트레스 저항성 조절과 관련하여, At5g21280 돌연변이체는 정상적인 생장조건에서는 잎의 크기나 유식물 생체량 등 바이오매스가 감소하는 표현형을 보이고 다른 생장과 발달에 표현형을 보이지 않지만 프로그램화 세포사멸을 유도하는 AAL-toxin, 캐탈라제 억제제인 아미노트리아졸 (aminotriazole)과 제초제의 일종인 파라콰트(paraquat)에 의해 유도되는 산화스트레스에 내성을 보이고, 이러한 산화스트레스 저항성은 비생물학적 스트레스 관련 전사인자 DREB19, HSFA2, ZAT10 등의 축적과 밀접한 연관이 있는 것으로 알려져 있다.Regarding vii) regulation of oxidative stress resistance in plants, the At5g21280 mutant shows a phenotype of decreased biomass such as leaf size and seedling biomass under normal growth conditions and shows no other growth and development phenotypes, but undergoes programmed cell death. It shows resistance to oxidative stress induced by AAL-toxin, aminotriazole, a catalase inhibitor, and paraquat, a type of herbicide, and this resistance to oxidative stress is due to the abiotic stress-related transcription factor DREB19. It is known to be closely related to the accumulation of , HSFA2, and ZAT10.
즉, 본 발명에 따른 sep 유전자는 i) 식물체의 지방산 함량 조절; ii) 식물체 내 글리옥시좀 또는 퍼옥시좀 발현 조절; iii) 식물세포의 크기 조절; iv) 식물체 내 지질 분해 조절; v) 식물체 내 글리옥실산 회로 관련 유전자 발현 조절; vi) 식물체의 개화시기 조절; 및 vii) 식물체의 산화스트레스 저항성 조절로 이루어지는 군으로부터 선택되는 어느 하나 이상의 특징을 가지고, 식물체 생장을 조절할 수 있다. 또한, 상기 "식물체 생장 조절"은 추가로 종자발아, 유식물 생중량, 하배축과 잎 크기, 개화시기를 조절하는 것을 특징으로써 가질 수 있다.That is, the sep gene according to the present invention i) regulates the fatty acid content of the plant; ii) regulation of glyoxisome or peroxisome expression in plants; iii) regulation of plant cell size; iv) regulation of lipid degradation in plants; v) Regulation of glyoxylic acid cycle-related gene expression in plants; vi) Control of flowering time of plants; and vii) regulating the oxidative stress resistance of the plant, and can control plant growth. In addition, the “plant growth control” can additionally be characterized by controlling seed germination, seedling fresh weight, hypocotyl and leaf size, and flowering time.
본 발명의 다른 하나의 양태는 sep 유전자를 포함하는 재조합 벡터를 제공한다.Another aspect of the present invention provides a recombinant vector containing a sep gene.
상기 재조합 벡터는 sep 유전자의 발현이 증가하거나, sep 유전자의 발현이 억제되도록 제작한 것일 수 있으나, 이에 제한되지 않는다. 상기 유전자의 발현을 증가 또는 억제시키는 벡터의 제조방법은 당업계에 알려져 있는 방법이면 제한 없이 사용할 수 있다.The recombinant vector may be designed to increase the expression of the sep gene or to suppress the expression of the sep gene, but is not limited thereto. Methods for producing vectors that increase or suppress the expression of the genes can be used without limitation as long as they are known in the art.
본 발명에서, 용어 "재조합"은 세포가 이종의 핵산을 복제하거나, 상기 핵산을 발현하거나 또는 펩티드, 이종의 펩티드 또는 이종의 핵산에 의해 코딩된 단백질을 발현하는 것을 지칭하는 것이다. 재조합 세포는 상기 세포의 천연 형태에서는 발견되지 않는 유전자 또는 유전자 단편을, 센스 또는 안티센스 형태 중 하나로 발현할 수 있다. 또한 재조합 세포는 천연 상태의 세포에서 발견되는 유전자를 발현할 수 있으며, 그러나 상기 유전자는 변형된 것으로서 인위적인 수단에 의해 세포 내 재도입된 것이다.In the present invention, the term “recombinant” refers to a cell replicating a heterologous nucleic acid, expressing the nucleic acid, or expressing a peptide, a heterologous peptide, or a protein encoded by a heterologous nucleic acid. Recombinant cells can express genes or gene fragments that are not found in the natural form of the cell, either in sense or antisense form. Additionally, recombinant cells can express genes found in cells in their natural state, but the genes have been modified and reintroduced into the cells by artificial means.
본 발명에서, 용어 "벡터"는 세포 내로 전달하는 DNA 단편 (들), 핵산 분자를 지칭할 때 사용된다. 벡터는 DNA를 복제시키고, 숙주세포에서 독립적으로 재생산될 수 있다. 용어 "전달체"는 흔히 "벡터"와 호환하여 사용된다. 용어 "발현 벡터"는 목적한 코딩 서열과, 특정 숙주 생물에서 작동가능하게 연결된 코딩 서열을 발현하는데 필수적인 적정 핵산 서열을 포함하는 재조합 DNA 분자를 의미한다. 진핵세포에서 이용 가능한 프로모터, 인핸서, 종결신호 및 폴리아데닐레이션 신호는 공지되어 있다.In the present invention, the term “vector” is used to refer to a DNA fragment(s) or nucleic acid molecule that is delivered into a cell. Vectors replicate DNA and can reproduce independently in host cells. The term “vector” is often used interchangeably with “vector”. The term “expression vector” refers to a recombinant DNA molecule containing a coding sequence of interest and an appropriate nucleic acid sequence necessary to express the operably linked coding sequence in a particular host organism. The promoters, enhancers, termination signals and polyadenylation signals available in eukaryotic cells are known.
본 발명의 벡터는 전형적으로 발현 또는 클로닝을 위한 벡터로서 구축될 수 있다. 또한, 본 발명의 벡터는 원핵세포 또는 진핵 세포를 숙주로 하여 구축될 수 있다. Vectors of the present invention can typically be constructed as vectors for expression or cloning. Additionally, the vector of the present invention can be constructed using prokaryotic cells or eukaryotic cells as hosts.
본 발명에서 용어 "프로모터"란 구조 유전자로부터의 DNA 업스트림의 영역을 의미하며 전사를 개시하기 위하여 RNA 중합효소가 결합하는 DNA 분자를 말한다. "식물 프로모터"는 식물 세포에서 전사를 개시할 수 있는 프로모터이다. 본 발명의 벡터에 포함되는 프로모터는 상기 조직 및 발달단계 비특이적 프로모터일 수 있고, 구체적으로 35S CaMV 프로모터일 수 있으나, 이에 제한되지 않는다.In the present invention, the term "promoter" refers to a region of DNA upstream from a structural gene and refers to a DNA molecule to which RNA polymerase binds to initiate transcription. A “plant promoter” is a promoter capable of initiating transcription in plant cells. The promoter included in the vector of the present invention may be a non-specific promoter for the tissue or developmental stage, and may specifically be the 35S CaMV promoter, but is not limited thereto.
본 발명의 재조합 벡터는 당업자에 주지된 방법에 의해 구축될 수 있다. 상기 방법은 시험관 내 재조합 DNA 기술, DNA 합성 기술 및 생체 내 재조합 기술 등을 포함한다. 상기 DNA 서열은 mRNA 합성을 이끌기 위해 발현 벡터 내의 적당한 프로모터에 효과적으로 연결될 수 있다. 또한 벡터는 번역 개시 부위로서 리보솜 결합 부위 및 전사 터미네이터를 포함할 수 있다.The recombinant vector of the present invention can be constructed by methods well known to those skilled in the art. The methods include in vitro recombinant DNA technology, DNA synthesis technology, and in vivo recombinant technology. The DNA sequence can be effectively linked to an appropriate promoter within an expression vector to drive mRNA synthesis. The vector may also include a ribosome binding site and a transcription terminator as a translation initiation site.
식물 발현 벡터의 바람직한 예는 아그로박테리움 튜머파시엔스 (Agrobacterium tumefiaciens)와 같은 적당한 숙주에 존재할 때 그 자체의 일부, 소위 T-영역을 식물 세포로 전이시킬 수 있는 Ti-플라스미드 벡터이다. 다른 유형의 Ti-플라스미드 벡터 (EP 0116718 B1 참조)는 현재 식물 세포, 또는 잡종 DNA를 식물의 게놈 내에 적당하게 삽입시키는 새로운 식물이 생산될 수 있는 원형질체로 잡종 DNA 서열을 전이시키는데 이용되고 있다. Ti-플라스미드 벡터의 특히 바람직한 형태는 EP 0120516 B1 및 미국 특허 제4,940,838호에 청구된 바와 같은 소위 바이너리 벡터이다. 본 발명에 따른 유전자를 식물 숙주에 도입시키는데 이용될 수 있는 다른 적합한 벡터는 이중가닥 식물 바이러스 (예를 들면, CaMV) 및 단일 가닥 바이러스, 게미니 바이러스 등 으로부터 유래될 수 있는 것과 같은 바이러스 벡터, 예를 들면 비완전성 식물 바이러스 벡터로부터 선택될 수 있다. 그러한 벡터의 사용은 특히 식물 숙주를 적당하게 형질전환 하는 것이 어려울 때 유리할 수 있다.A preferred example of a plant expression vector is the Ti-plasmid vector, which is capable of transferring part of itself, the so-called T-region, into plant cells when present in a suitable host such as Agrobacterium tumefaciens . Another type of Ti-plasmid vector (see EP 0116718 B1) is currently used to transfer hybrid DNA sequences into plant cells or protoplasts from which new plants can be produced that properly integrate the hybrid DNA into the plant's genome. Particularly preferred forms of Ti-plasmid vectors are the so-called binary vectors as claimed in EP 0120516 B1 and US Pat. No. 4,940,838. Other suitable vectors that can be used to introduce genes according to the invention into plant hosts include viral vectors, such as those that may be derived from double-stranded plant viruses (e.g., CaMV) and single-stranded viruses, geminiviruses, etc. For example, it may be selected from non-intact plant virus vectors. The use of such vectors can be particularly advantageous when it is difficult to properly transform plant hosts.
발현 벡터는 바람직하게는 하나 이상의 선택성 마커를 포함한다. 상기 마커는 통상적으로 화학적인 방법으로 선택될 수 있는 특성을 갖는 핵산 서열로, 형질전환된 세포를 비형질전환 세포로부터 구별할 수 있는 모든 유전자가 이에 해당된다. 상기 마커 유전자는 항생제 내성 유전자 (antibiotics resistance gene)일 수 있으나, 이에 제한되지 않는다.The expression vector preferably contains one or more selectable markers. The marker is a nucleic acid sequence that has characteristics that can be generally selected by chemical methods, and includes all genes that can distinguish transformed cells from non-transformed cells. The marker gene may be an antibiotic resistance gene, but is not limited thereto.
본 발명의 일 구현 예에 따른 식물 발현 벡터에서, 터미네이터는 통상의 터미네이터를 사용할 수 있으며, 그 예로는 노팔린 신타아제 (NOS), 벼 α-아밀라아제 RAmy1 A 터미네이터, 파세올린 (phaseoline) 터미네이터, 아그로박테리움 튜머파시엔스 (Agrobacterium tumefaciens)의 옥토파인 합성 (Octopine synthase) 유전자의 터미네이터 등이 있으나, 이에 제한되지 않는다.In the plant expression vector according to one embodiment of the present invention, the terminator may be a conventional terminator, examples of which include nopaline synthase (NOS), rice α-amylase RAmy1 A terminator, phaseoline terminator, Agro Examples include, but are not limited to, the terminator of the Octopine synthase gene of Agrobacterium tumefaciens .
본 발명의 벡터를 진핵 세포에 형질전환시키는 경우에는 숙주세포로서, 효모 (Saccharomyce cerevisiae), 곤충세포, 사람세포 (예컨대, CHO 세포주 (Chinese hamster ovary), W138, BHK, COS-7, 293, HepG2, 3T3, RIN 및 MDCK 세포주) 및 식물세포 등이 이용될 수 있다. 본 발명에 있어서, 숙주세포는 구체적으로 식물세포일 수 있다.When the vector of the present invention is transformed into eukaryotic cells, yeast ( Saccharomyce cerevisiae ), insect cells, human cells (e.g., CHO cell line (Chinese hamster ovary), W138, BHK, COS-7, 293, HepG2 are used as host cells. , 3T3, RIN and MDCK cell lines) and plant cells can be used. In the present invention, the host cell may specifically be a plant cell.
본 발명의 벡터를 숙주세포 내로 운반하는 방법은, 숙주 세포가 원핵 세포인 경우, CaCl2 방법, 하나한 방법 (Hanahan D. Studies on transformation of Escherichia coli with plasmids. Journal of Molecular Biology. 1983;166(4):557-580) 및 전기천공 방법 등에 의해 실시될 수 있다. 또한, 숙주세포가 진핵세포인 경우에는, 미세주입법, 칼슘포스페이트 침전법, 전기천공법, 리포좀-매개 형질감염법, DEAE-덱스트란 처리법, 및 유전자 밤바드먼트 등에 의해 벡터를 숙주세포 내로 주입할 수 있으나, 이에 제한되지 않는다.The method of transporting the vector of the present invention into a host cell is, when the host cell is a prokaryotic cell, the CaCl 2 method, the Hanahan method (Hanahan D. Studies on transformation of Escherichia coli with plasmids. Journal of Molecular Biology. 1983;166 ( 4):557-580) and electroporation methods. In addition, when the host cell is a eukaryotic cell, the vector can be injected into the host cell by microinjection, calcium phosphate precipitation, electroporation, liposome-mediated transfection, DEAE-dextran treatment, and gene bombardment. may, but is not limited to this.
본 발명의 다른 하나의 양태는 식물체 내에서 sep 유전자를 과발현시키는 단계를 포함하는, 식물체의 생장 촉진 방법을 제공한다.Another aspect of the present invention provides a method for promoting plant growth, comprising the step of overexpressing the sep gene in the plant.
상기 sep 유전자의 과발현은 식물세포에 sep 유전자를 포함하는 재조합 벡터를 운반하여 식물체 내에서 sep 유전자가 과발현 되도록 유도하는 것일 수 있으나, 이에 제한되지 않는다. 상기 sep 유전자를 포함하는 재조합 벡터 및 이를 숙주세포인 식물세포 내로 운반하는 방법 등에 대해서는 전술한 바와 같다.Overexpression of the sep gene may be carried out by delivering a recombinant vector containing the sep gene to a plant cell to induce overexpression of the sep gene in the plant, but is not limited to this. The recombinant vector containing the sep gene and the method of transporting it into a plant cell as a host cell are as described above.
상기 sep 유전자는 서열번호 1 또는 서열번호 2의 염기서열로 표시되는 것일 수 있고, 상기 sep 유전자는 서열번호 3 또는 서열번호 4의 아미노산 서열을 코딩하는 것일 수 있다. 이에 대해서는 전술한 바와 같다.The sep gene may be represented by the base sequence of SEQ ID NO: 1 or SEQ ID NO: 2, and the sep gene may encode the amino acid sequence of SEQ ID NO: 3 or SEQ ID NO: 4. This is the same as described above.
상기 sep 유전자가 과발현되는 대상 식물체는 구체적으로 애기장대, 담배, 가지, 고추, 토마토, 우엉, 쑥갓, 상추, 도라지, 시금치, 근대, 고구마, 샐러리, 당근, 미나리, 파슬리, 배추, 양배추, 갯무, 수박, 참외, 오이 호박, 박, 딸기, 대두, 녹두, 강낭콩, 및 완두인 쌍자엽 식물 또는 벼, 보리, 밀, 호밀, 옥수수, 사탕수수, 귀리, 및 양파 등의 단자엽 식물일 수 있고, 보다 구체적으로 쌍자엽 식물일 수 있으며, 더욱 구체적으로 애기장대일 수 있으나, 이에 제한되지 않는다.Target plants overexpressing the sep gene are specifically Arabidopsis thaliana, tobacco, eggplant, pepper, tomato, burdock, mugwort, lettuce, bellflower root, spinach, Swiss chard, sweet potato, celery, carrot, water parsley, parsley, Chinese cabbage, cabbage, radish, It may be a dicotyledonous plant such as watermelon, melon, cucumber pumpkin, gourd, strawberry, soybean, mung bean, kidney bean, and pea, or a monocotyledonous plant such as rice, barley, wheat, rye, corn, sugarcane, oats, and onion, and may be more specific. It may be a dicotyledonous plant, and more specifically, it may be Arabidopsis thaliana, but is not limited thereto.
상기 식물체 생장 촉진은 다음 중 선택되는 어느 하나 이상의 특징을 가지는 것인, 식물체의 생장 촉진 방법:A method of promoting plant growth, wherein the plant growth promotion has one or more characteristics selected from the following:
i) 식물체의 지방산 함량 감소;i) reducing the fatty acid content of the plant;
ii) 식물체 내 글리옥시좀 또는 퍼옥시좀 발현 증가;ii) increased expression of glyoxysomes or peroxisomes in plants;
iii) 식물세포의 크기 증가;iii) increase in size of plant cells;
iv) 식물체 내 지질 분해 증가;iv) increased lipid breakdown in the plant;
v) 식물체 내 글리옥실산 회로 관련 유전자 발현 증가;v) increased expression of glyoxylic acid cycle-related genes in plants;
vi) 식물체의 개화시기 촉진; 및vi) Promoting the flowering time of plants; and
vii) 식물체의 산화스트레스 저항성 증가.vii) Increased resistance to oxidative stress in plants.
전술한 본 발명의 구현예서는 상기 sep 유전자에 변이가 있는 경우 식물체가 잘 자라지 못하여 생장 및 개화시기가 지연되는 것을 확인하고, 상기 sep 유전자에 변이가 있는 식물체에 sep 유전자를 과발현시키는 경우, 야생형 수준으로 정상적으로 생장 및 개화하는 것을 확인하였는 바, 이에 따라 식물체에서 sep 유전자를 과발현할 경우 i) 식물체의 지방산 함량 감소; ii) 식물체 내 글리옥시좀 또는 퍼옥시좀 발현 증가; iii) 식물세포의 크기 증가; iv) 식물체 내 지질 분해 조절; v) 식물체 내 글리옥실산 회로 관련 유전자 발현 증가; vi) 식물체의 개화시기 증가; 및 vii) 식물체의 산화스트레스 저항성 증가로 이루어지는 군으로부터 선택되는 어느 하나 이상의 특징을 가지고, 식물체 생장을 촉진할 수 있다. In the above-described embodiment of the present invention, it is confirmed that when there is a mutation in the sep gene, the plant does not grow well and the growth and flowering time are delayed, and when the sep gene is overexpressed in a plant with a mutation in the sep gene, the plant grows at the wild-type level. It was confirmed that the plant grows and blooms normally. Accordingly, when the sep gene is overexpressed in the plant, i) the fatty acid content of the plant is reduced; ii) increased expression of glyoxysomes or peroxisomes in plants; iii) increase in size of plant cells; iv) regulation of lipid degradation in plants; v) increased expression of glyoxylic acid cycle-related genes in plants; vi) Increased flowering time of plants; and vii) increasing the oxidative stress resistance of the plant, and can promote plant growth.
또한, 상기 "식물체 생장 성장"은 추가로 종자발아를 촉진하고, 유식물 생중량, 하배축과 잎 크기를 증가시키는 것을 특징으로써 가질 수 있다.In addition, the “plant growth” can additionally be characterized by promoting seed germination and increasing seedling fresh weight, hypocotyl and leaf size.
본 발명의 다른 하나의 양태는 식물체 내에서 sep 유전자의 발현을 억제시키는 단계를 포함하는, 식물체의 생장 저해 방법을 제공한다.Another aspect of the present invention provides a method of inhibiting the growth of a plant, comprising the step of suppressing the expression of the sep gene in the plant.
상기 sep 유전자의 발현 억제는 식물세포에 sep 유전자를 억제하도록 제작한 재조합 벡터를 운반하여 식물체 내에서 sep 유전자의 발현이 억제되도록 유도하는 것일 수 있으나, 이에 제한되지 않는다. 상기 sep 유전자를 억제하도록 제작한 재조합 벡터 및 이를 숙주세포인 식물세포 내로 운반하는 방법 등에 대해서는 전술한 바와 같다.Suppression of the expression of the sep gene may be carried out in plant cells by delivering a recombinant vector designed to suppress the sep gene, thereby inducing the expression of the sep gene to be suppressed in the plant, but is not limited to this. The recombinant vector designed to suppress the sep gene and the method of transporting it into a plant cell as a host cell are as described above.
상기 sep 유전자는 서열번호 1 또는 서열번호 2의 염기서열로 표시되는 것일 수 있고, 상기 sep 유전자는 서열번호 3 또는 서열번호 4의 아미노산 서열을 코딩하는 것일 수 있다. 이에 대해서는 전술한 바와 같다.The sep gene may be represented by the base sequence of SEQ ID NO: 1 or SEQ ID NO: 2, and the sep gene may encode the amino acid sequence of SEQ ID NO: 3 or SEQ ID NO: 4. This is the same as described above.
상기 sep 유전자 발현이 억제되는 대상 식물체는 구체적으로 애기장대, 담배, 가지, 고추, 토마토, 우엉, 쑥갓, 상추, 도라지, 시금치, 근대, 고구마, 샐러리, 당근, 미나리, 파슬리, 배추, 양배추, 갯무, 수박, 참외, 오이 호박, 박, 딸기, 대두, 녹두, 강낭콩, 및 완두인 쌍자엽 식물 또는 벼, 보리, 밀, 호밀, 옥수수, 사탕수수, 귀리, 및 양파 등의 단자엽 식물일 수 있고, 보다 구체적으로 쌍자엽 식물일 수 있으며, 더욱 구체적으로 애기장대일 수 있으나, 이에 제한되지 않는다.Target plants in which sep gene expression is suppressed are specifically Arabidopsis thaliana, tobacco, eggplant, pepper, tomato, burdock, mugwort, lettuce, bellflower root, spinach, Swiss chard, sweet potato, celery, carrot, water parsley, parsley, Chinese cabbage, cabbage, and radish. , watermelons, melons, cucumbers, pumpkins, gourds, strawberries, soybeans, mung beans, kidney beans, and peas, or monocots such as rice, barley, wheat, rye, corn, sugarcane, oats, and onions. Specifically, it may be a dicotyledonous plant, and more specifically, it may be Arabidopsis thaliana, but is not limited thereto.
상기 식물체 생장 저해는 다음 중 선택되는 어느 하나 이상의 특징을 가지는 것인, 식물체의 생장 저해 방법:A method of inhibiting plant growth, wherein the plant growth inhibition has one or more characteristics selected from the following:
i) 식물체의 지방산 함량 증가;i) increasing the fatty acid content of the plant;
ii) 식물체 내 글리옥시좀 또는 퍼옥시좀 발현 감소;ii) Reduced expression of glyoxysomes or peroxisomes in plants;
iii) 식물세포의 크기 감소;iii) Reduction in size of plant cells;
iv) 식물체 내 지질 분해 감소;iv) reduction of lipid breakdown in plants;
v) 식물체 내 글리옥실산 회로 관련 유전자 발현 감소;v) Reduced expression of genes related to the glyoxylic acid cycle in plants;
vi) 식물체의 개화시기 지연; 및vi) delay in flowering of plants; and
vii) 식물체의 산화스트레스 저항성 감소.vii) Reduction of oxidative stress resistance in plants.
전술한 본 발명의 구현예서는 상기 sep 유전자에 변이가 있는 경우 식물체가 잘 자라지 못하여 생장 및 개화시기가 지연되는 것을 확인하였는 바, 이에 따라 식물체에서 sep 유전자의 발현을 억제할 경우 i) 식물체의 지방산 함량 증가; ii) 식물체 내 글리옥시좀 또는 퍼옥시좀 발현 감소; iii) 식물세포의 크기 감소; iv) 식물체 내 지질 분해 감소; v) 식물체 내 글리옥실산 회로 관련 유전자 발현 감소; vi) 식물체의 개화시기 지연; 및 vii) 식물체의 산화스트레스 저항성 감소 로 이루어지는 군으로부터 선택되는 어느 하나 이상의 특징을 가지고, 식물체 생장을 억제할 수 있다. In the above-described embodiment of the present invention, it was confirmed that when there is a mutation in the sep gene, the plant does not grow well and the growth and flowering time are delayed. Accordingly, when the expression of the sep gene is suppressed in the plant, i) the fatty acid of the plant increased content; ii) Reduced expression of glyoxysomes or peroxisomes in plants; iii) Reduction in size of plant cells; iv) reduction of lipid breakdown in plants; v) Reduced expression of genes related to the glyoxylic acid cycle in plants; vi) delay in flowering of plants; and vii) reducing the oxidative stress resistance of the plant, and can inhibit plant growth.
또한, 상기 "식물체 생장 저해"는 추가로 종자발아를 지연시키고, 유식물 생중량, 하배축과 잎 크기를 감소시키는 것을 특징으로써 가질 수 있다.In addition, the “plant growth inhibition” may additionally be characterized by delaying seed germination and reducing seedling fresh weight, hypocotyl and leaf size.
본 발명의 다른 하나의 양태는 본 발명의 재조합 벡터로 형질전환된 식물체를 제공한다. Another aspect of the present invention provides a plant transformed with the recombinant vector of the present invention.
본 발명의 다른 하나의 양태는 본 발명의 재조합 벡터로 형질전환된 식물체의 종자를 제공한다. Another aspect of the present invention provides seeds of plants transformed with the recombinant vector of the present invention.
상기 재조합 벡터에 대해서는 전술한 바와 같다.The recombinant vector is as described above.
상기 식물체는 애기장대, 담배, 가지, 고추, 토마토, 우엉, 쑥갓, 상추, 도라지, 시금치, 근대, 고구마, 샐러리, 당근, 미나리, 파슬리, 배추, 양배추, 갯무, 수박, 참외, 오이 호박, 박, 딸기, 대두, 녹두, 강낭콩, 완두, 벼, 보리, 밀, 호밀, 옥수수, 사탕수수, 귀리, 및 양파로 이루어지는 군으로부터 선택되는 어느 하나 이상일 수 있고, 구체적으로 애기장대일 수 있으나, 이에 제한되지 않는다.The plants include Arabidopsis thaliana, tobacco, eggplant, pepper, tomato, burdock, mugwort, lettuce, bellflower root, spinach, chard, sweet potato, celery, carrot, water parsley, parsley, Chinese cabbage, cabbage, radish, watermelon, melon, cucumber, pumpkin, and gourd. , strawberries, soybeans, mung beans, kidney beans, peas, rice, barley, wheat, rye, corn, sugar cane, oats, and onions, and may specifically be Arabidopsis thaliana, but are not limited thereto. No.
본 발명의 또 하나의 양태는 본 발명의 재조합 벡터로 식물세포를 형질전환하는 단계; 및 상기 형질전환된 식물세포로부터 식물을 재분화하는 단계;를 포함하는 식물체 생장이 조절된 형질전환 식물체의 제조방법을 제공한다. Another aspect of the present invention includes the steps of transforming plant cells with the recombinant vector of the present invention; and redifferentiating plants from the transformed plant cells. A method for producing a transformed plant with controlled plant growth is provided.
상기 재조합 벡터에 대해서는 전술한 바와 같다.The recombinant vector is as described above.
상기 제조방법은 본 발명의 재조합 벡터로 식물세포를 형질전환하는 단계를 포함하며, 식물세포로의 형질전환은 DNA를 식물세포에 전이시키는 임의의 방법을 사용할 수 있다. 식물 종의 형질전환은 쌍자엽 식물뿐만 아니라 단자엽 식물 양자를 포함한 식물 종에 대해 일반적이다. 원칙적으로, 임의의 형질전환 방법은 본 발명에 따른 잡종 DNA를 적당한 선조 세포로 도입시키는데 이용될 수 있다. 상기 방법은 원형질체에 대한 칼슘/폴리에틸렌 글리콜 방법, 원형질체의 전기천공법, 식물 요소로의 현미주사법, 각종 식물 요소의 (DNA 또는 RNA-코팅된) 입자 충격법, 식물의 침윤 또는 성숙 화분 또는 소포자의 형질전환에 의한 아그로박테리움 튜머파시엔스 매개된 유전자 전이에서 (비완전성) 바이러스에 의한 감염 등으로부터 적당하게 선택될 수 있다. 그러한 형질전환 방법은 반드시 재생 및(또는) 조직 배양기간을 가질 필요는 없다.The production method includes the step of transforming plant cells with the recombinant vector of the present invention, and transformation into plant cells can be performed using any method for transferring DNA to plant cells. Transformation of plant species is common for plant species including both monocots as well as dicots. In principle, any transformation method can be used to introduce the hybrid DNA according to the invention into suitable progenitor cells. The methods include the calcium/polyethylene glycol method for protoplasts, electroporation of protoplasts, microinjection into plant elements, particle bombardment (DNA or RNA-coated) of various plant elements, infiltration of plants or maturation of pollen or spores. It can be appropriately selected from Agrobacterium tumefaciens mediated gene transfer by transformation, infection by a (non-complete) virus, etc. Such transformation methods do not necessarily require a regeneration and/or tissue culture period.
또한, 본 발명의 제조방법은 상기 형질전환된 식물 세포로부터 형질전환 식물을 재분화하는 단계를 포함한다. 형질전환 식물 세포로부터 형질전환 식물을 재분화하는 방법은 당업계에 공지된 임의의 방법을 이용할 수 있다.Additionally, the production method of the present invention includes the step of redifferentiating the transformed plant from the transformed plant cells. A method for redifferentiating a transgenic plant from a transgenic plant cell can use any method known in the art.
본 발명에 따라 조직/발달단계 비특이적 프로모터와 종자 식물 특이 익스텐신 모티브를 포함하는 단백질 (seeds plant specific extensin motif containing proteins, SEP) 코딩 서열이 연결된 재조합 벡터를 사용하여 지질분해, 글리옥실산 회로, 글리옥시좀/퍼옥시좀 발생 조절이 가능하므로, 식물의 유지 함량과 조성을 제어를 통한 유지식물 및 유지작물의 개량 또는 개발에 유용하게 활용될 수 있다.According to the present invention, lipolysis, glyoxylic acid cycle, Since the generation of glyoxysomes/peroxisomes can be controlled, it can be usefully used in the improvement or development of oil plants and oil crops by controlling the oil content and composition of the plant.
도 1a는 종자 식물 특이 익스텐신 모티브를 포함하는 단백질 (seeds plant specific extensin motif containing proteins, SEP) 동원체(ortholog)의 계통수 분석 결과이다. 단자엽 클레이드가 애기장대를 포함하는 쌍자엽 클레이드와 나누어진다.
도 1b는 애기장대와 밀 및 대표적인 작물 (A.thaliana: 애기장대 (Arabidopsis thaliana), C.sativa: 삼 (Cannabis sativa), B.rapa: 배추 (Brassica rapa), T.hassleriana: 풍접초 (Tarenaya hassleriana), J.regia: 가래나무 (Juglans regia), N.nucifera: 연꽃 (Nelumbo nucifera), T.aestivum: 밀 (Triticum aestivum), O.sativa: 벼 (Oryza sativa))의 SEP 동원체의 다서열 배열을 시도한 결과이다. 핵으로 이동하는 서열 NLS가 매우 잘 보존되어 있다.
도 2a는 TAIR로부터 분양받은 애기장대 Salk_006796 라인이 엑손1에 T-DNA가 삽입위치를 RT-PCR로 결정한 결과이다. 2개의 액손과 1개의 인트론으로 구성되어 있다. 인트론이 제거되고 엑손 1과 2가 연결된 변이체 1 (variant 1, Chr5:7263828..7265874)과 엑손1 염기서열 말단에 위치하는 종료코돈에 의해서 엑손 1만 발현되는 변이체 2 (variant 2, Chr5:7263880..7265851) 두 종류의 유전자 모델을 갖고 있다 (https://www.arabidopsis.org/servlets/TairObject?name=AT5G21280&type=locus).
도 2b는 서열번호 1 및 서열번호 2의 아미노산 서열을 암호화하는 서열번호 3 및 서열번호 4가 연결된 재조합 벡터를 나타낸 모식도이다. 프로모터는 35S CaMV를 사용하여 조직과 발달단계 비특이적으로 발현시켰으며, 세포내 위치와 항원항체 반응을 위해 GFP 태깅하였다.
도 2c는 애기장대 원형질체에서 GFP 표진된 AtSEP가 DAPI가 염색되는 세포내 위치인 핵에 존재함을 확인하였다.
도 2d는 호모와 헤테로 보상 발현체 계통에서 AtSEP가 발현됨을 GFP를 항원항체 반으로 확인하였다.
도 3은 유식물 (a)과 영양생장 (b) 및 생식생장기 (c)의 야생형 애기장대 식물체 (Col-O), Atsep 엑손 1에 T-DNA가 삽입된 돌연변이체 Salk_006796 (sep), Salk_006796의 유전적 배경에 애기장대 Atsep를 과발현시킨 보상발현체 (AtCO)의 표현형 (하배축 길이, 잎 면적 및 개화시기)를 조사한 결과이다.
도 4a와 b는 상기 도 3a에서 관찰된 유식물 하배축을 대상으로 세포길이와 lipid body를 Nile red 염색약으로 관찰한 후 지방산 대사물을 정량한 결과이다. 도 4c는 상기 식물체를 대상으로 포도당 처리에 의해 sep 돌연변이 식물체의 유식물 생장이 회복됨을 확인한 결과이다.
도 5는 상기 도 4에서 관찰된 유식물을 대상으로 전사체 발현 분석을 하여 지방분해 경로와 글리옥시좀/퍼옥시좀 생성에 관여하는 유전자 발현을 야생종과 비교하여 상대적으로 계산한 결과이다.
도 6은 상기 도 3의 조건에서 sep 돌연변이체 유전적 배경에 밀 sep 유전자를 보상발현한 식물체 (TaCO)에서 하배축 길이, 잎 면적, 개화시기 및 표피세포 길이가 야생종 수준으로 부분 혹은 완전히 회복됨을 보여주는 결과이다.Figure 1a shows the results of phylogenetic tree analysis of orthologs of seeds plant specific extensin motif containing proteins (SEP). The monocot clade is divided from the dicot clade, which includes Arabidopsis thaliana.
Figure 1b shows Arabidopsis thaliana, wheat and representative crops (A.thaliana: Arabidopsis thaliana ), C.sativa: Hemp ( Cannabis sativa ), B.rapa: Chinese cabbage ( Brassica rapa ), T.hassleriana: Tarenaya hassleriana ), J.regia: Juglans regia , N.nucifera: lotus ( Nelumbo nucifera ), T.aestivum: wheat ( Triticum aestivum ), O.sativa: multi-sequence arrangement of the SEP centromere of rice ( Oryza sativa ) This is the result of trying. The sequence NLS that migrates to the nucleus is highly conserved.
Figure 2a shows the results of determining the location of T-DNA insertion in exon 1 of the Arabidopsis Salk_006796 line purchased from TAIR by RT-PCR. It consists of two axons and one intron. Variant 1 (variant 1, Chr5:7263828..7265874), in which the intron is removed and exons 1 and 2 are connected, and variant 2 (variant 2, Chr5:7263880), in which only exon 1 is expressed by a stop codon located at the end of the exon 1 base sequence. ..7265851) It has two types of gene models (https://www.arabidopsis.org/servlets/TairObject?name=AT5G21280&type=locus).
Figure 2b is a schematic diagram showing a recombinant vector in which SEQ ID NO: 3 and SEQ ID NO: 4 encoding the amino acid sequences of SEQ ID NO: 1 and SEQ ID NO: 2 are linked. The promoter was expressed tissue- and development-stage non-specifically using 35S CaMV, and was tagged with GFP for intracellular location and antigen-antibody reaction.
Figure 2c confirmed that GFP-labeled AtSEP was present in the nucleus, an intracellular location where DAPI was stained, in Arabidopsis protoplasts.
Figure 2d shows that AtSEP is expressed in homo and hetero compensatory expression lines, confirming the GFP antigen-antibody assay.
Figure 3 shows wild-type Arabidopsis plants (Col-O) in seedling (a), vegetative growth (b), and reproductive growth (c), and mutants Salk_006796 ( sep ) and Salk_006796 with T-DNA inserted into Atsep exon 1. This is the result of examining the phenotype (hypocotyl length, leaf area, and flowering time) of a compensatory expression construct (AtCO) that overexpressed Arabidopsis Atsep in a genetic background.
Figures 4a and b show the results of quantifying fatty acid metabolites after observing cell length and lipid body with Nile red dye in the seedling hypocotyl observed in Figure 3a. Figure 4c shows the results confirming that seedling growth of the sep mutant plant was restored by glucose treatment of the plant.
Figure 5 shows the results of transcript expression analysis for the seedlings observed in Figure 4 above and calculating relative expression of genes involved in the lipolysis pathway and glyoxysome/peroxisome production compared to wild species.
Figure 6 shows that hypocotyl length, leaf area, flowering time, and epidermal cell length are partially or completely restored to the level of wild species in plants (TaCO) that compensated for the expression of the wheat sep gene in the sep mutant genetic background under the conditions of Figure 3. It is a result.
이하, 하기 실시예에 의하여 본 발명을 보다 상세하게 설명한다. 단, 하기 실시예는 본 발명을 예시하기 위한 것일 뿐 본 발명의 범위가 이들로 한정되는 것은 아니다.Hereinafter, the present invention will be described in more detail through the following examples. However, the following examples are only for illustrating the present invention and the scope of the present invention is not limited thereto.
실시예 1. 애기장대 및 밀의 익스텐신Example 1. Extensins from Arabidopsis thaliana and wheat (( extensin, EXTextensin, EXT )) 모티브 분석 Motif analysis
EXT 모티브는 세린 1개와 연이어 3~5개의 프롤린으로 구성된 펩티드 (SerPro3~5)로서, 애기장대의 경우 177개의 유전자가 알려져 있다. 이중 42개는 SerPro3~5 모티브 주위에 Tyr-Val-Tyr 모티브를 갖는 익스텐신이다. 11개는 익스텐신 모티브 외에 Leu과 Cys이 반복되는 도메인 (LRR)을 포함하는데 이를 LRR-EXT라 한다. 이외에 아라비노갈락탄 (arabinogalactan) 단백질-EXT (4개), PERK (15개), 포르민 (액신-미세소관 결합도메인, 11개)가 있다. 이들은 세포신장, 신호감지 및 전달, 세포골격 형태조절에 관여하는 것으로 알려져 있다 (Borassi, Cecilia & Sede, Ana & Mecchia, Martin & Salter, Juan & Marzol, Eliana & Muschietti, Jorge & Estevez, Josι. (2015). An update on cell surface proteins containing extensin-motifs. J. Exp. Bot. 67).The EXT motif is a peptide (SerPro3~5) composed of one serine followed by three to five prolines, and in Arabidopsis thaliana, 177 genes are known. Of these, 42 are extensins with Tyr-Val-Tyr motifs around the SerPro3 to 5 motifs. In addition to the extensin motif, 11 contain a Leu and Cys repeating domain (LRR), which is called LRR-EXT. In addition, there are arabinogalactan proteins - EXT (4), PERK (15), and formin (axin-microtubule binding domain, 11). They are known to be involved in cell elongation, signal sensing and transmission, and regulation of cytoskeletal morphology (Borassi, Cecilia & Sede, Ana & Mecchia, Martin & Salter, Juan & Marzol, Eliana & Muschietti, Jorge & Estevez, Josι. (2015 ). An update on cell surface proteins containing extensin-motifs. J. Exp. Bot. 67).
애기장대 At5G21280 유전자위 (atr7 유전자)가 코딩하는 단백질 또한 2개의 익스텐신 모티브를 갖는 단백질 (서열번호 3)이지만 상기 서술한 5가지의 익스텐신형 세포표면 단백질에는 포함되지 않는다. The protein encoded by the Arabidopsis At5G21280 locus (atr7 gene) is also a protein with two extensin motifs (SEQ ID NO: 3), but is not included in the five extensin-type cell surface proteins described above.
이에, SeqNLS 분석(http://mleg.cse.sc.edu/seqNLS/) 결과 애기장대 At5G21280 유전자위가 코딩하는 단백질은 핵 수송 서열 (KKQVRRR)을 포함하고 있어 세포외로 분비되는 세포표면 단백질에 포함되지 않을 것이라는 견해를 뒷받침한다. 애기장대 At5G21280 유전자위에 해당하는 서열은 애기장대 이외에 다수의 쌍자엽 식물과 밀을 포함하는 단자엽 식물에서만 발견되고 조류나 하등식물에서는 발견되지 않는다 (도 1a). Accordingly, as a result of SeqNLS analysis (http://mleg.cse.sc.edu/seqNLS/), the protein encoded by the Arabidopsis thaliana At5G21280 locus contains a nuclear transport sequence (KKQVRRR) and is therefore included in the cell surface proteins secreted extracellularly. This supports the view that this will not happen. The sequence corresponding to the Arabidopsis thaliana At5G21280 locus is found only in many dicots other than Arabidopsis and monocots, including wheat, and is not found in algae or lower plants (Figure 1a).
다서열배열(multiple sequence analysis, MSA) 분석 결과 애기장대 At5G21280 유전자위가 코딩하는 단백질 내 두 영역 (KQVRRRLHTSRPYQERLLNMAEARREIVTALK (서열번호 16) 및 LNFLLPNQPLGLNLNFQDFNDFiQT (서열번호 17))이 애기장대를 포함하는 다양한 식물 (A.thaliana: 애기장대 (Arabidopsis thaliana), C.sativa: 삼 (Cannabis sativa), B.rapa: 배추 (Brassica rapa), T.hassleriana: 풍접초 (Tarenaya hassleriana), J.regia: 가래나무 (Juglans regia), N.nucifera: 연꽃 (Nelumbo nucifera), T.aestivum: 밀 (Triticum aestivum), O.sativa: 벼 (Oryza sativa)) 내에서 잘 보존되어 있음을 확인하였다 (도 1b). As a result of multiple sequence analysis (MSA) analysis, two regions (KQVRRRLHTSRPYQERLLNMAEARREIVTALK (SEQ ID NO: 16) and LNFLLPNQPLGLNLNFQDFNDFiQT (SEQ ID NO: 17)) within the protein encoded by the Arabidopsis thaliana At5G21280 locus were found in various plants (A. thaliana: Arabidopsis thaliana , C.sativa: Hemp ( Cannabis sativa ), B.rapa: Chinese cabbage ( Brassica rapa ), T.hassleriana: Tarenaya hassleriana , J.regia: Juglans regia , It was confirmed that it was well preserved in N.nucifera: lotus ( Nelumbo nucifera ), T.aestivum: wheat ( Triticum aestivum ), and O.sativa: rice ( Oryza sativa ) (Figure 1b).
애기장대 At5G21280 유전자위가 코딩하는 단백질을 2차 구조 예측 프로그램으로 분석한 결과 막통과 나선 (transmembrane helices)이 없는 수용성 단백질인 것을 알 수 있으나, Pfam 도메인 서치를 통해서는 특정한 도메인이 예측되지 않았으며, 특정 3차원 구조 분석 (https://iupred2a.elte.hu/)에서도 예측되지 않는 본질적으로 구조화되지 않은 단백질 (Intrinsically unstructured protein)으로 분류되었다. As a result of analyzing the protein encoded by the Arabidopsis thaliana At5G21280 locus using a secondary structure prediction program, it was found to be a water-soluble protein without transmembrane helices, but no specific domain was predicted through Pfam domain search. It was classified as an intrinsically unstructured protein that was not predicted by specific 3D structure analysis (https://iupred2a.elte.hu/).
한편, OrthoDB v10.1 (https://www.orthodb.org/) 분석 결과에서는 애기장대 At5G21280 유전자위가 코딩하는 단백질에서는 프롤린 잔기가 다수 발견되어 하이드록시 프롤린이 풍부한 당단백질 계열 단백질 (hydroxyproline-rich glycoprotein family protein)로도 분류되지만, 프롤린 풍부 단백질 (Pro-rich Protein, PRP)의 특징인 펜타펩타이드 모티프 (pentapeptide motif, PPVX(K/T))를 포함하고 있지 않기 때문에 이미 4종이 알려져 있는 애기장대 PRP에는 포함되지 않는다. Meanwhile, in the results of OrthoDB v10.1 (https://www.orthodb.org/) analysis, many proline residues were found in the protein encoded by the Arabidopsis thaliana At5G21280 locus, indicating that it is a hydroxyproline-rich glycoprotein family protein (hydroxyproline-rich). Although it is classified as a glycoprotein family protein, it does not contain the pentapeptide motif (PPVX(K/T)), which is a characteristic of proline-rich protein (PRP), so there are already four known types of Arabidopsis PRP. is not included in
상기 결과를 바탕으로 이하에서 애기장대 At5G21280 유전자위가 코딩하는 단백질은 애기장대 유래 종자 식물 특이 익스텐신 모티브를 포함하는 단백질 (seeds plant specific extensin motif containing proteins, SEP), 즉, AtSEP로 명명하고, 애기장대 At5G21280 유전자위는 애기장대 유래 sep 으로 명명하였다. Based on the above results, hereinafter, the protein encoded by the Arabidopsis thaliana At5G21280 locus is named Arabidopsis-derived seed plant specific extensin motif containing proteins (SEP), that is, AtSEP, The Arabidopsis thaliana At5G21280 gene locus was named sep from Arabidopsis thaliana.
한편, EnsemblPlants (http://plants.ensembl.org/index.html)에서 AtSEP 아미노산 서열을 퀘리로 BLASTP를 수행하여 밀 (Triticum aestivum)로부터 선발된 6종 단백질 (> 60% 동일성) 중에서 AtSEP와 가장 큰 아미노산 서열 동일성을 보이는 단백질 (서열번호 4)을 선별하고, 이를 코딩하는 유전자인 TraesCS5B02G518900의 염기서열 (서열번호 2)을 갖는 DNA를 합성하였다 (Bioneer). 상기 AtSEP와 아미노산 서열 동일성이 큰 단백질 및 이를 코딩하는 유전자인 TraesCS5B02G518900.1을 각각 밀 유래 SEP (TaSEP) 및 sep로 명명하였다.Meanwhile, BLASTP was performed by querying the AtSEP amino acid sequence in EnsemblPlants (http://plants.ensembl.org/index.html), and among the six proteins (>60% identity) selected from wheat ( Triticum aestivum ), AtSEP was the most A protein showing large amino acid sequence identity (SEQ ID NO: 4) was selected, and DNA having the nucleotide sequence (SEQ ID NO: 2) of the gene encoding it, TraesCS5B02G518900, was synthesized (Bioneer). The protein with high amino acid sequence identity to AtSEP and the gene encoding it, TraesCS5B02G518900.1, were named wheat-derived SEP (TaSEP) and sep, respectively.
실시예 2. Salk_006796 라인의 T-DNA 삽입 위치 및 변이체 1과 2의 발현량 분석Example 2. Analysis of T-DNA insertion location and expression level of variants 1 and 2 in Salk_006796 line
ABRC (Arabidopsis Biological Resource Center, ABRC)에서 At5g21280 유전자에 대한 T-DNA 삽입 돌연변이체(Salk_006796, Salk_081956, 및 cs820583) 3종을 제작하고, 이중 엑손1이 삽입되어 있는 Salk_006796을 실험에 사용하였다 (도 2a). At ABRC (Arabidopsis Biological Resource Center, ABRC), three T-DNA insertion mutants (Salk_006796, Salk_081956, and cs820583) for the At5g21280 gene were created, and Salk_006796, which has exon 1 inserted, was used in the experiment (Figure 2a) ).
0067LP (서열번호 5)와 0067RP (서열번호 6) 프라이머로 RT-PCR한 결과, 애기장대 야생종 (Col-0)에서는 밴드가 관찰되었으나 Salk_006796 돌연변이체에서는 밴드가 관찰되지 않았다. 반면 T-DNA 상의 프라이머 LBb1.3 (서열번호 7)과 0067R과의 RT-PCR에서는 Salk_006796 돌연변이체에서만 밴드가 관찰되었다 (도 2a).As a result of RT-PCR using primers 0067LP (SEQ ID NO: 5) and 0067RP (SEQ ID NO: 6), a band was observed in the wild type of Arabidopsis thaliana (Col-0), but no band was observed in the Salk_006796 mutant. On the other hand, in RT-PCR with primers LBb1.3 (SEQ ID NO: 7) and 0067R on T-DNA, a band was observed only in the Salk_006796 mutant (FIG. 2a).
sep 전장과 3'UTR을 포함하는 프라이머 (서열번호 8 및 서열번호 9)를 이용한 RT-PCR 결과에서 Col-0에서는 sep cDNA 밴드가 관찰되었으나 Salk-007696 돌연변이체에서는 관찰되지 않았다. 이때 PCR 산물은 분자량 크기와 서열 분석 결과 변이체 1 (variant 1)에 해당하였다 (도 2a). In RT-PCR results using primers containing the full length of sep and 3'UTR (SEQ ID NO: 8 and SEQ ID NO: 9), a sep cDNA band was observed in Col-0, but not in the Salk-007696 mutant. At this time, the PCR product corresponded to variant 1 as a result of molecular weight and sequence analysis (Figure 2a).
실시예 3. 애기장대 SEP 또는 밀 유래 SEP 코딩 서열을 포함하는 벡터 제작 Example 3. Construction of a vector containing the coding sequence of Arabidopsis SEP or wheat-derived SEP
35S CaMV 프로모터가 결합된 애기장대 SEP 또는 밀 SEP 코딩 서열을 각각 포함하는 재조합 벡터인 35SCsMV:AtSEP:GFP 또는 35SCsMV:TaSEP:GFP를 제작하여 애기장대 SEP 돌연변이체 (SALK_006796)에 형질전환하였다. 35SCsMV:AtSEP:GFP or 35SCsMV:TaSEP:GFP, a recombinant vector containing the Arabidopsis SEP or wheat SEP coding sequence, respectively, combined with the 35S CaMV promoter, was constructed and transformed into the Arabidopsis SEP mutant (SALK_006796).
애기장대의 경우, Col-0 에서 식물 총 RNA 추출 키트 (SpectrumTM Plant Total RNA extraction kit, Sigma-Aldrich)를 이용하여 RNA를 500 ng 추출한 뒤, 이를 cDNA 합성 키트 (iScriptTM cDNA synthesis kit, Bio-rad)를 이용하여 cDNA로 합성하였다. AtSEP 스플라이싱 변이체 1에 해당하는 cds를 PCR로 증폭한 뒤, AtSEP에 GFP (Green Fluorescent Protein), myc, flag을 표지하여 CaMV35S 프로모터로 발현시킬 수 있는 JJ461 벡터에 클로닝하였다. In the case of Arabidopsis thaliana, 500 ng RNA was extracted from Col-0 using a plant total RNA extraction kit (Spectrum TM Plant Total RNA extraction kit, Sigma-Aldrich), and then extracted with a cDNA synthesis kit (iScript TM cDNA synthesis kit, Bio- rad) was used to synthesize cDNA. The cds corresponding to AtSEP splicing variant 1 was amplified by PCR, and then AtSEP was labeled with GFP (Green Fluorescent Protein), myc, and flag and cloned into the JJ461 vector that can be expressed with the CaMV35S promoter.
밀의 경우, TaSEP에 GFP를 표지하여 CaMV35S 프로모터로 발현시킬 수 있는 JJ461 벡터에 클로닝하였다 (도 2b). 여기에서 사용되는 프라이머 서열은 하기 표 1과 같다.In the case of wheat, TaSEP was labeled with GFP and cloned into the JJ461 vector that can be expressed using the CaMV35S promoter (Figure 2b). The primer sequences used here are shown in Table 1 below.
(서열번호 12)AATCTAGAATGAGGAATCAAAGATTAGCT
(SEQ ID NO: 12)
(서열번호 13)AAGGATCCACGATAACCACTCCTCATC
(SEQ ID NO: 13)
실시예 4. 애기장대 형질전환 식물체 제작 Example 4. Production of Arabidopsis transgenic plants
실시예 3에서 벡터에 클로닝한 유전자는 아그로박테리움 GV3101에 형질전환 후 플로럴 딥 방법 (floral dip method (Clough and Bent, 1998))을 이용하여 애기장대에 도입하였다. 형질전환된 애기장대는 항생제를 함유한 1/2 MS 배지(1/2 MS, 1% 수크로스, 20 ug/ml 하이그로마이신 (hygromycin), 1% 피토아가 (phytoagar), pH 5.8) 에서 3:1로 분리된 라인을 선별 후, T3 이상의 동형접합 식물체를 실험에 사용하였다. 식물 배양시 종자 표면은 70% EtOH로 5분, 0.8% 차아염소산 나트륨 (sodium hypochlorite)으로 10분간 살균한 후, 증류수로 5회 세척하였다. 종자 휴면을 깨기 위해 48시간 동안 암실에서 저온 처리 (4℃)하였다. 배양실 온도는 22℃, 16/8 시간의 광주기, 빛 세기는 100 μmol m-2s-1 이었다. 실험에 사용한 모든 종자는 수확한 후 22℃에서 2주간 후숙하여 사용하였다. The gene cloned into the vector in Example 3 was transformed into Agrobacterium GV3101 and then introduced into Arabidopsis thaliana using the floral dip method (Clough and Bent, 1998). Transformed Arabidopsis thaliana was grown in 1/2 MS medium (1/2 MS, 1% sucrose, 20 ug/ml hygromycin, 1% phytoagar, pH 5.8) containing antibiotics for 3 days. After selecting the lines separated by :1, homozygous plants of T3 or higher were used in the experiment. During plant culture, the seed surface was sterilized with 70% EtOH for 5 minutes, 0.8% sodium hypochlorite for 10 minutes, and then washed 5 times with distilled water. To break seed dormancy, the seeds were treated at low temperature (4°C) in the dark for 48 hours. The culture room temperature was 22°C, a photoperiod of 16/8 hours, and light intensity was 100 μmol m -2 s -1 . All seeds used in the experiment were harvested and post-ripened for 2 weeks at 22°C.
전사량이 확인된 ATSEP (2 라인)와 TaSEP (3 라인) 형질전환체의 5 라인을 GFP 단클론 항체를 이용하여 웨스턴 블롯 분석을 수행하였다. 구체적으로, 7일 자란 식물체에서 단백질을 추출하였으며, 50 ㎍ 단백질을 8% SDS-PAGE 젤에서 분리하여 PVDF (Polyvinylidene fluoride) 막 (Merck, Darmstadt, Germany)에 옮기고 GFP 단클론 항체 (Sigma, St. Louis, MO, USA), 염소 항 마우스 IgG-HRP (horseradish peroxidase) (Santa Cruz Biotechnology, Inc., Santa Cruz, CA, USA) 및 웨스턴 ECL 기질 (BIO-RAD clarity Max western ECL substrate, Bio-Rad, Hercules, CA, USA)을 사용하여 웨스턴 블롯 분석하였다. Western blot analysis was performed on 5 lines of ATSEP (line 2) and TaSEP (line 3) transformants with confirmed transcription amounts using a GFP monoclonal antibody. Specifically, proteins were extracted from plants grown for 7 days, and 50 μg of protein was separated on an 8% SDS-PAGE gel, transferred to PVDF (Polyvinylidene fluoride) membrane (Merck, Darmstadt, Germany), and GFP monoclonal antibody (Sigma, St. Louis). , MO, USA), goat anti-mouse IgG-HRP (horseradish peroxidase) (Santa Cruz Biotechnology, Inc., Santa Cruz, CA, USA) and western ECL substrate (BIO-RAD clarity Max western ECL substrate, Bio-Rad, Hercules , CA, USA) was used for Western blot analysis.
그 결과, 형질전환 식물체에서 SEP 단백질은 실시예 1에서의 핵 수송 모티브 존재의 확인 및 기존에 보고된 바 (Sujeeth, Neerakkal et al. (2019). A novel seed plants gene regulates oxidative stress tolerance in Arabidopsis thaliana. Cellular and Molecular Life Sciences. 77))와 같이 핵에 위치하는 것을 확인하였다 (도 2d). As a result, the SEP protein in the transgenic plant confirmed the presence of a nuclear transport motif in Example 1 and was previously reported (Sujeeth, Neerakkal et al. (2019). A novel seed plants gene regulates oxidative stress tolerance in Arabidopsis thaliana Cellular and Molecular Life Sciences. 77)) was confirmed to be located in the nucleus (Figure 2d).
실시예 5. 애기장대 sep 돌연변이 식물체의 표현형 분석Example 5. Phenotypic analysis of Arabidopsis sep mutant plants
실시예 3의 벡터로 형질전환된어 보상발현된 sep 돌연변이 식물체의 분자유전학적 기능을 확인하기 위해, Col-0, 이전 연구에서 제작된 p296 (Dong Ho Shin, et al., Identification of genes that may regulate the expression of the transcription factor production of anthocyanin pigment 1 (PAP1)/MYB75 involved in Arabidopsis anthocyanin biosynthesis, Plant Cell Rep (2015) 34:805-815.), 실시예 2의 Salk_006796 (sep) 및 실시예 3의 벡터로 형질전환된어 보상발현된 sep 돌연변이 식물체 (AtCO)를 22℃, 16/8 시간의 광주기 조건에서 종자발아, 유식물, 영양생장 및 생식생장 전 발달 단계의 표현형을 분석하였다. To confirm the molecular genetic function of sep mutant plants transformed with the vector of Example 3 and compensated for expression, Col-0 and p296 produced in a previous study (Dong Ho Shin, et al., Identification of genes that may regulate the expression of the transcription factor production of anthocyanin pigment 1 (PAP1)/MYB75 involved in Arabidopsis anthocyanin biosynthesis, Plant Cell Rep (2015) 34:805-815.), Salk_006796 ( sep ) of Example 2 and Example 3 The vector-transformed sep mutant plants (AtCO) with compensatory expression were analyzed for the phenotypes of seed germination, seedlings, vegetative growth, and pre-reproductive growth stages at 22°C and a photoperiod of 16/8 hours.
야생형을 비롯한 sep 돌연변이체(p296, sep)과 보상발현된 sep 돌연변이 식물체 (AtCO)의 유식물 (5일 생육한 식물체, 도 3a)에서 영양생장 (19일 생육한 식물체, 도 3b) 및 생식생장 단계(37일 생육한 식물체, 도 3c))의 sep 발현에 따른 발달 및 생장속도를 관찰한 결과, Salk_006796 (sep)는 5일 생육한 유식물의 뿌리 및 식물체의 크기와 19일 생육한 식물체의 자엽 및 rosette 잎의 크기가 Salk_006796 (sep)가 Col-0에 비해 작았으며, 37일 생육한 식물체에서 꽃대 및 꽃의 발달이 야생형에 비해 지연되었다. 이러한 발아 및 생장 지연 표현형은 sep 도입에 의해 야생형 수준으로 회복되었다 (AtCO). Vegetative growth (plants grown on 19 days, Figure 3b) and reproductive growth in seedlings (plants grown on 5 days, Figure 3a) of sep mutants ( p296 , sep ), including the wild type, and sep mutant plants with compensatory expression (AtCO). As a result of observing the development and growth rate according to sep expression at the stage (plants grown on 37 days, Figure 3c)), Salk_006796 ( sep ) was the root and plant size of seedlings grown on 5 days and the size of plants grown on 19 days. The size of cotyledons and rosette leaves in Salk_006796 ( sep ) was smaller than that in Col-0, and the development of flower stalks and flowers in plants grown for 37 days was delayed compared to the wild type. These germination and growth retardation phenotypes were restored to wild-type levels by introduction of sep (AtCO).
상기 결과는 조직 혹은 발단 단계에서 sep 유전자에 의해 식물체의 종자 발아 지연과 왜소성 및 개화 조절이 가능함을 시사한다. The above results suggest that it is possible to delay seed germination, dwarfism, and control flowering in plants by the sep gene at the tissue or germination stage.
실시예 6. 애기장대 SEP의 세포길이 생장 조절 기전 분석Example 6. Analysis of the cell length growth regulation mechanism of Arabidopsis SEP
식물길이는 세포의 분열 (division)과 신장 (elongation)에 의해서 결정된다. 실시예 5에서 sep 돌연변이체인 Salk_006796에서 확인된 생장 억제는 세포분열보다 세포신장 억제에서 기인하는 것으로 예상되었다. 이를 확인하기 위해, 1/2 MS, 1.5% 피토아가 함유 배지에서 온도와 빛 조건을 달리하여 7일 생육한 Col-0, Salk_006796 (sep) 및 실시예 3의 벡터로 형질전환된어 보상발현된 sep 돌연변이 식물체 (AtCO)의 유식물 하배축 세포크기를 측정하였다. Plant length is determined by cell division and elongation. The growth inhibition confirmed in the sep mutant Salk_006796 in Example 5 was expected to result from inhibition of cell elongation rather than cell division. To confirm this, Col-0, Salk_006796 ( sep ) grown for 7 days in a medium containing 1/2 MS and 1.5% phytoaga under different temperature and light conditions, and were transformed with the vector of Example 3 and compensated for expression. Seedling hypocotyl cell size of sep mutant plants (AtCO) was measured.
그 결과, Salk_006796 (sep)는 Col-0에 비해서 하배축 표피 장축 길이가 54% 수준으로 감소하였으나 AtCO에서는 77-80% 수준의 길이를 보였다 (도 4a 아래). 상기 하배축 표피 세포 길이 관찰 시 sep 돌연변이 식물체에서 물방울 모양의 액체성 물체가 관측되었는데, Nile Red로 염색되는 유적 (oil droplet)으로 확인되었다 (도 4a 위). As a result, Salk_006796 ( sep ) had a hypocotyl epidermal long axis length reduced to 54% compared to Col-0, but AtCO showed a length of 77-80% (bottom of Figure 4a). When observing the hypocotyl epidermal cell length, water drop-shaped liquid objects were observed in the sep mutant plants, which were confirmed to be oil droplets stained with Nile Red (Figure 4a above).
상기 배양조건에서 7일 생육한 Col-0, p296, Salk_006796 (sep) 및 AtCO의 유식물 하배축을 0.0025% 나일 레드 (Nile Red, N3013, Sigma-aldrich) 용액으로 37℃에서 10분간 처리한 후 공초점현미경(LSM800, CarlZeiss)으로 관찰한 결과, Col-0과 AtCO에서는 소량 형광이 관찰된 반면 p296과 Salk_006796 (sep)에서는 현저히 많은 유적이 관찰되었다 (도 4b). 암소에서 7일 생장되었을 경우에도 유사한 형광분포를 보였다. The seedling hypocotyls of Col-0, p296 , Salk_006796 ( sep ) and AtCO grown for 7 days under the above culture conditions were treated with 0.0025% Nile Red (N3013, Sigma-aldrich) solution at 37°C for 10 minutes, then incubated. As a result of observation using a focal microscope (LSM800, CarlZeiss), a small amount of fluorescence was observed in Col-0 and AtCO, while significantly more traces were observed in p296 and Salk_006796 ( sep ) (Figure 4b). A similar fluorescence distribution was observed when grown in the dark for 7 days.
생육한 Col-0, Salk_006796 (sep) 및 AtCO 내 지방산 함량 및 구성 성분을 밝히고자 가스크로마토그래피로 분석하였다. 동결 건조된 종자 및 암조건에서 7일 생육한 유식물 10 mg을 2.5% 황산과 톨루엔을 넣고 90℃에서 1시간 30분간 처리하였다. 1M NaCl과 헥산을 넣고 원심분리하여 상층액 900ul 분리하였고, 이를 화염 이온화 검출기 (Flame Ionization Detector) 와 INNOWAX 모세관 컬럼 (INNOWAX capillary column, 30 m x 0.32 mm x 0.5 um, Agilent Techonologies)을 이용하여 가스크로마토크래피 (YL-6100GC, YoungLin Science) 분석에 사용하였다. 표준시료로 펜타데칸산 (C15:0) 10 mg을 사용하였다 (Meesapyodsuk, Dauenpen & Qiu, Xiao. (2008). An Oleate Hydroxylase from the Fungus Claviceps purpurea: Cloning, Functional Analysis, and Expression in Arabidopsis. Plant physiology. 147. 1325-33; Siloto, Rodrig, et al. (2006). The Accumulation of Oleosins Determines the Size of Seed Oilbodies in Arabidopsis. The Plant cell. 18. 1961-74). To reveal the fatty acid content and composition in grown Col-0, Salk_006796 ( sep ) and AtCO, they were analyzed by gas chromatography. 10 mg of freeze-dried seeds and seedlings grown for 7 days under dark conditions were treated with 2.5% sulfuric acid and toluene at 90°C for 1 hour and 30 minutes. 1M NaCl and hexane were added and centrifuged to separate 900ul of the supernatant, which was subjected to gas chromatography using a flame ionization detector and INNOWAX capillary column (30 mx 0.32 mm x 0.5 um, Agilent Technologies). RAPY (YL-6100GC, YoungLin Science) was used for analysis. 10 mg of pentadecanoic acid (C15:0) was used as a standard sample (Meesapyodsuk, Dauenpen & Qiu, Xiao. (2008). An Oleate Hydroxylase from the Fungus Claviceps purpurea: Cloning, Functional Analysis, and Expression in Arabidopsis. Plant physiology 147. 1325-33; Siloto, Rodrig, et al. (2006). The Accumulation of Oleosins Determines the Size of Seed Oilbodies in Arabidopsis. The Plant cell. 18. 1961-74).
그 결과, 암조건에서 7일 생육한 Salk_006796 (sep)의 총 지방산 함량은 Col-0에 비해 약 377% 높았으며, AtCO의 총 지방산 함량은 Col-0와 유사하였다 (도 4b). As a result, the total fatty acid content of Salk_006796 ( sep ) grown for 7 days in dark conditions was about 377% higher than Col-0, and the total fatty acid content of AtCO was similar to Col-0 (Figure 4b).
종자에 저장된 지질은 글리옥시좀/퍼옥시좀이 관여하는 분해대사 경로를 통해서 하배축을 포함하는 생장축 (growing axis)에서 필요로 하는 탄소원과 에너지원으로 사용된다. 이에, Salk_006796 (sep) 돌연변이 식물에서 유적의 축적에 의해서 포도당 신생성이 억제되고, 결과적으로 하배축 등 유식물이 억제되었을 경우에 외부에서 탄소원을 공급하면, 돌연변이체 생장이 야생종 수준으로 회복될 것으로 예상되었다. 1/2 MS, 1.5% 피토아가에 각 1%의 단당류와 이당류를 처리하였다 (Glu: 포도당, Fru: 과당, Suc: 수크로스, Mal: 말토오스). 삼투압에 대한 대조군으로 만니톨 (Man)과 소비톨 (Sor)을 처리하였다. Lipids stored in seeds are used as carbon and energy sources required by the growing axis, including the hypocotyl, through a metabolic metabolic pathway involving glyoxysomes/peroxisomes. Accordingly, in Salk_006796 ( sep ) mutant plants, gluconeogenesis is suppressed by the accumulation of remains, and as a result, when seedlings such as hypocotyls are suppressed, it is expected that the growth of the mutant will recover to the level of wild species if carbon sources are supplied from outside. It has been done. 1/2 MS, 1.5% phytoaga was treated with 1% each of monosaccharides and disaccharides (Glu: glucose, Fru: fructose, Suc: sucrose, Mal: maltose). Mannitol (Man) and sorbitol (Sor) were treated as controls for osmotic pressure.
그 결과, 대조군 탄소원을 처리한 배지에서 7일 생장한 Salk_006796 (sep)의 생중량은 야생형의 26% 수준이었으나 포도당 처리한 배지에서 Salk_006796 (sep)의 생중량은 Col-0 대비 각각 90% 수준으로 회복되었다. 그러나 과당, 이당류, 오스모티쿰인 소비톨과 만니톨 처리에 의해서는 이러한 생장 회복현상이 나타나지 않았다 (도 4c). As a result, the fresh weight of Salk_006796 ( sep ) grown for 7 days in the medium treated with the control carbon source was 26% of the wild type, but the fresh weight of Salk_006796 ( sep ) in the medium treated with glucose was 90% of that of Col-0. recovered. However, this growth recovery phenomenon did not occur when treated with fructose, disaccharide, osmoticum sorbitol, and mannitol (Figure 4c).
상기 결과는 SEP가 저장 지질 분해에 관여하는 것을 나타내고, 나아가 sep 발현을 조절함으로써 지방산 분해 의존성 에너지 및 탄소대사를 통해서 세포길이 조절이 가능함을 시사한다.The above results indicate that SEP is involved in the breakdown of storage lipids, and further suggest that controlling sep expression allows for cell length control through fatty acid breakdown-dependent energy and carbon metabolism.
실시예 7. 애기장대 SEP의 지질 대사 유전자 발현 조절 분석Example 7. Analysis of lipid metabolism gene expression regulation of Arabidopsis SEP
TAG (triacylglycerol)와 같은 종자 저장 지질이나 엽록체에서 생성된 지방산은 글리옥시좀/퍼옥시좀의 베타-산화경로에 의해서 아세틸 코에이 (aceteyl-CoA) 로 분해된다. 아세틸 코에이는 글리옥실산 회로, 포도당 신생성 경로와 시트르산 회로를 통해서 메발론산 경로 등 유식물 생장에 필요한 탄소골격 전구체와 에너지를 생성하는 것으로 알려져 잇다. 잎 퍼옥시좀에 의한 베타-산화경로는 지방산 수준을 일정하게 유지할 뿐만 아니라 일시적인 녹말 결핍 조건에서 대사 항상성을 유지하는 역할을 하는 것으로 알려져 있다. 베타산화 경로 유전자 돌연변이 식물체에서 볼 수 있는 유식물 생장 저해나 지방산 축적에 의한 엽록체 손상이 외부에서 공급된 포도당에 의해서 억제된다는 결과는 지질 분해 대사의 중요성을 시사한다. Seed storage lipids such as TAG (triacylglycerol) or fatty acids produced in chloroplasts are decomposed into acetyl-CoA by the beta-oxidation pathway of glyoxysomes/peroxisomes. Acetyl CoA is known to generate carbon skeleton precursors and energy necessary for seedling growth, including the mevalonic acid pathway, through the glyoxylic acid cycle, gluconeogenic pathway, and citric acid cycle. The beta-oxidation pathway by leaf peroxisomes is known to not only maintain constant fatty acid levels but also plays a role in maintaining metabolic homeostasis under temporary starch deficiency conditions. The results showing that the inhibition of seedling growth and chloroplast damage due to fatty acid accumulation seen in beta-oxidation pathway gene mutant plants are suppressed by externally supplied glucose suggest the importance of lipolytic metabolism.
즉, 도 3에서 확인된 Salk_006796 (sep)에서 세포길이 생장 저해는 유적의 집적과 연관되어 있으며, 도 4에서 확인된 외부에서 공급된 포도당에 의해 표현형이 야생형인 Col-0 수준으로 복구되는 결과는 SEP의 작용표적이 글리옥시좀/퍼옥시좀에 의해서 매개되는 지방산 분해 경로임을 시사한다. 특히, SEP가 핵 소재 단백질 시그널을 포함하고 있는 핵 단백질이므로 (도 1b, 도 2d), 퍼옥시좀이 관여하는 지방산 분해 경로에 관여할 것으로 예상되었다. That is, the inhibition of cell length growth in Salk_006796 ( sep ) identified in Figure 3 is associated with the accumulation of remnants, and the result of restoring the phenotype to the level of wild-type Col-0 by externally supplied glucose confirmed in Figure 4 This suggests that the target of action of SEP is the fatty acid degradation pathway mediated by glyoxysomes/peroxisomes. In particular, since SEP is a nuclear protein containing a nuclear localization protein signal (Figure 1b, Figure 2d), it was expected to be involved in the fatty acid degradation pathway involving peroxisomes.
이에, 퍼옥시좀이 관여하는 지방산 분해 경로 유전자 발현을 분석하였다. 발아 직후 (2일 째) 및 7일 생장된 식물에서 실시예 3의 방법으로 총 RNA를 추출한 후, mRNA 샘플 준비 키트 (Illumina TruSeq Stranded mRNA Sample Preparation kit, Illumina) 로 mRNA 라이브러리를 제작하였다. 이어서 RNA 시퀀스 분석 (NextSeq500 sequencers, Illumina)을 수행한 결과, 예상한 바와 같이 Salk_006796 (sep)에서 TAG 모빌라이제이션 (SDP1), 베타 산화 (PXA1, ACX2-4, KAT2), 글리옥실산 회로 (CSY3, ACO3, MLS, PMDH1) 그리고 퍼옥시좀 생성 (PEX11A, D, E)과 관련된 유전자량이 야생종에 비해서 2~8배 정도 감소하였다. 이러한 유전자량 감소는 AtCO에서는 Col-0 수준으로 회복되었다 (도 5). Accordingly, the expression of genes in the fatty acid degradation pathway involving peroxisomes was analyzed. Total RNA was extracted from plants immediately after germination (day 2) and grown for 7 days by the method of Example 3, and then an mRNA library was prepared using an mRNA sample preparation kit (Illumina TruSeq Stranded mRNA Sample Preparation kit, Illumina). Subsequent RNA sequence analysis (NextSeq500 sequencers, Illumina) showed that, as expected, TAG mobilization (SDP1), beta oxidation (PXA1, ACX2-4, KAT2), and glyoxylate cycle (CSY3) were detected in Salk_006796 ( sep ). , ACO3, MLS, PMDH1) and peroxisome production (PEX11A, D, E) decreased by 2 to 8 times compared to the wild species. This reduction in gene mass was restored to the Col-0 level in AtCO (Figure 5).
실시예 8. 밀 SEP에 의한 애기장대 sep 돌연변이 표현형의 야생형으로의 회복Example 8. Recovery of Arabidopsis sep mutant phenotype to wild type by wheat SEP
밀 SEP는 애기장대 SEP와 달리 단자엽 클레이드에 포함되며, 잘 보존되어 있는 도메인 사이에 변이가 매우 크다 (도 1). 이러한 서열상의 변이에도 불구하고 실시예 3의 밀 SEP 벡터를 이용하여 애기장대 sep 돌연변이체에 밀 SEP를 이형발현 (heterologus expression) 시켰을 경우 하배축 길이와 세포길이와 잎의 길이 및 개화시기가 부분 혹은 온전히 회복되었다 (도 6). Unlike Arabidopsis thaliana SEPs, wheat SEPs are included in the monocot clade, and there is great variation among well-conserved domains (Figure 1). Despite these sequence variations, when wheat SEP was heterologously expressed in an Arabidopsis sep mutant using the wheat SEP vector of Example 3, the hypocotyl length, cell length, leaf length, and flowering time were partially or completely changed. recovered (Figure 6).
상기 결과는 변이 지역보다는 보존서열 영역이 SEP 생화학적 기능에 중요함을 시사하였다.The above results suggested that the conserved sequence region rather than the mutation region is important for SEP biochemical function.
상기 실시예의 결과로부터, 애기장대 또는 밀 유래의 핵 단백질 SEP가 지방산 분해와 퍼옥시좀 생성에 관여하는 유전자량을 조절함으로써 식물의 생장과 개화시기 등을 조절하는 것을 확인하였으며, sep 돌연변이 시 지방산 분해대사가 저해되어 식물체 하배축, 잎 면적, 생체량, 개화시기, 세포길이 생장이 지연되는 바, sep 유전자를 포함하는 재조합 벡터로 식물세포를 형질전환시켜 sep 유전자의 발현을 조절함으로써, 식물체의 지방산 함량과 조성, 생장과 개화시기, 종자발아, 식물세포 크기, 식물체 내 지질분해, 식물체 내 글리옥실산 회로, 및 글리옥시좀/퍼옥시좀 생산을 조절할 수 있다.From the results of the above examples, it was confirmed that SEP, a nuclear protein derived from Arabidopsis thaliana or wheat, regulates plant growth and flowering time by controlling the amount of genes involved in fatty acid decomposition and peroxisome production, and fatty acid decomposition during sep mutation. As metabolism is inhibited, the growth of plant hypocotyl, leaf area, biomass, flowering time, and cell length is delayed. By transforming plant cells with a recombinant vector containing the sep gene and controlling the expression of the sep gene, the fatty acid content of the plant and Composition, growth and flowering time, seed germination, plant cell size, lipid decomposition in the plant, glyoxylic acid cycle in the plant, and glyoxysome/peroxisome production can be controlled.
이상의 설명으로부터, 본 발명이 속하는 기술분야의 당업자는 본 발명이 그 기술적 사상이나 필수적 특징을 변경하지 않고서 다른 구체적인 형태로 실시될 수 있다는 것을 이해할 수 있을 것이다. 이와 관련하여, 이상에서 기술한 실시 예들은 모든 면에서 예시적인 것이며 한정적인 것이 아닌 것으로서 이해해야만 한다. 본 발명의 범위는 상기 상세한 설명보다는 후술하는 특허 청구범위의 의미 및 범위 그리고 그 등가 개념으로부터 도출되는 모든 변경 또는 변형된 형태가 본 발명의 범위에 포함되는 것으로 해석되어야 한다.From the above description, those skilled in the art to which the present invention pertains will understand that the present invention can be implemented in other specific forms without changing its technical idea or essential features. In this regard, the embodiments described above should be understood in all respects as illustrative and not restrictive. The scope of the present invention should be construed as including the meaning and scope of the patent claims described below rather than the detailed description above, and all changes or modified forms derived from the equivalent concept thereof are included in the scope of the present invention.
<110> REPUBLIC OF KOREA(MANAGEMENT : RURAL DEVELOPMENT ADMINISTRATION) <120> Plant growth regulatory genes encoding seeds plant specific extensin motif containing proteins and uses thereof <130> KPA200896-KR <160> 17 <170> KoPatentIn 3.0 <210> 1 <211> 2047 <212> DNA <213> Unknown <220> <223> At5G21280 nucleotide <400> 1 caaaagtaca cacagagaca ctgttagctc ttaagtaata tgtcatcgta agcaacaaaa 60 aaaaatgaaa gagcaactaa ttctcatttc tgataatatt ttctgatttt tcacaaaaat 120 aaaattctgt aaaattgggc tagagcagag tatttatatc acccacgagc ctttgcttct 180 tcttccttct tcatcacctc tcttcttctt tctgttctct ttccggagac atttgtctaa 240 atctcatcat cattcacaat ttactcaaac aaaaccttga gaaagagaaa aaaatgatcc 300 aagagaagaa gaaaaaccta ttaatgtctt ctcagtgaaa aaccataact aaaatcttta 360 atggtcagat caaacaagca agaaccacgc gtctcttcta cttacatccg atctctcgtg 420 aagcaacaac ttgcttattc caccactatg accaccacca ccacaacaac aacaaacgat 480 ggtagcggcg gcggaaaaac gcaaacgcaa acgcacaaga aacaagtgag gaggcgactt 540 cacacaagcc gtccttatca agaacgtctc ctaaacatgg ctgaagctcg tcgcgaaatc 600 gtcaccgctt taaaacaaca tcgtgcttcc atgagacaag ccacaagaat cccaccgcct 660 caaccacccc caccaccgca gcctctaaat cttttctctc cgccgcctcc tcctcctcct 720 ccggatccat tttcttggac gaatccatct ctcaatttcc tcctccctaa tcaaccctta 780 gggttaaacc taaacttcca agatttcaac gacttcatcc aaacctcctc aacaacatct 840 tcatcatcat cttcctccac ttcatcttcc tcatcctcga tttttccgac gaatcctcat 900 atctactctt ccccttcacc tcctcccact ttcaccaccg ccacttcaga ttcagctcct 960 caactaccgt cttcctctaa tggagaaaac aacgtggtga cgtcagcttg gtggtcagaa 1020 ctcatgttga agacggtgga gccggagatt aaaccggaaa cggaagaagt catcgtagtc 1080 gaagatgacg tgtttccaaa gtttagtgac gtcatggaat ttccttcgtg gttaaaccaa 1140 acagaggaag aattatttca tccttataat ctcactgatc attactcatc atctcctcat 1200 aatcctccat tatcctggta attattttgt atttattttt cattacttag aattaaagtt 1260 attttgatcg gtgttgtttt tcctccggaa tcaccggtgg cgaaaattgc ggcggttctt 1320 gcataaactt cagtgattga gtttgagaca gaacacagaa aaaacagaat cgttcttttg 1380 tttggttttg ataattaaat cggagttatt ggattaaaaa tatgtgaaag cttgtggtat 1440 gagaaggatc cacataggtt tattgcacga tctgaagctg tccatttttt tcttttagta 1500 tatggtccaa tgataaataa tatcgccgac atctataaaa cactatcctt accccatttt 1560 tctgaactgt cccaaaaccg tcttgacgtc acccgatata aattatattt ttgttatgtt 1620 atcattgcac ttgcattatt caaatttggt tgttacttgt taggtaaatt atatatctct 1680 tttgtgtgtt ctaatgaaac atttttgatg tgtgatattc tttttacagt atggagattg 1740 gagaaattga aggcatggat ggagacgact ggcttgcttg aaccacaaag attgattact 1800 gttttttttt tcagacatga tgaatcaact ctttggttat tttcttatta tttttacctt 1860 attttatttg tttttcttta taattttttc tttttgtttg gtgacatttt aatgaatctt 1920 taaaaataac aacaagcctg cggagagaaa atgttcgatt tggcaaagcc acatgaatga 1980 agtaaaactt tcattttcgt tgattgactt tgttttttgt taataactta ttattagact 2040 ttaaatg 2047 <210> 2 <211> 1155 <212> DNA <213> Unknown <220> <223> TraesCS5B02G518900.1 nucleotide <400> 2 atgaggaacc agaggctagc tcctatcact tgctcctctt caaagaccaa caacaccaag 60 cgagaggagc cacacctctc cggggcttac attaggagcc ttgtgaagca tcttagctcc 120 tcatctacgg cgagatccaa agaccaccat cacatcacca tgggctccaa atcacaccaa 180 gaagagcaac aagccccaca aaccacacca tcatctctac agcaacagca gccacacaag 240 aagcaggtga ggaggaggct ccacacgagc aggccatatc aggagaggct actcaacatg 300 gcggaggcca ggcgagagat cgtcaccgct ctcaagatcc acagagcctc catgagagaa 360 gccaaggagc agcagcagca tcaacaactt gtgcaacaat tgcagcatca gcaagaagtc 420 caggtagtgc aagatcatag ggtagctttt agtgcaccca acatgagctc atatggttcc 480 ttctcagatt acttgcacaa tagttaccca ttcgaacact ccacagcacc aagcaacagt 540 ggttgctcat attattcatc tcctccactc ctcccttacc acacaccagt cgttgcaccc 600 atggttccca tggtagatgc tttagatcag ttgctgtcgc tgcctacgca gccactaggg 660 ctcaacctga cattcgatgg cttcaatggt ggtgttgctg ccgaagatgc caagaactgc 720 actgctacta gcccctttga tcctccttcg ttggtccaac aaccatcacc tgcctcctcc 780 tactctgtct actcctctcc accgctggcg acgatggtga gccaggacat ggcgtccgct 840 gctgccgaga acacctcaca gttgctgcac cgggtgctgg acgaggagga gatggcggcg 900 atccactcgg ccggggagcg gcatgacatc gagtggagcg acaccgtgaa cctggccacg 960 tcggcgtggt ggagcaggct actcgagagt gtggaaggcg gtgaagacga tgatggggcc 1020 gcggcggcgc agcagactaa cacggtggac gccatgggga tgcatctgag tgatgagtgc 1080 tacggccaag acgtgtcctt tccatgtatg gacatcggag agatcgaagg atgggacgcc 1140 gagtggttct catga 1155 <210> 3 <211> 302 <212> PRT <213> Unknown <220> <223> At5G21280 amino acid <400> 3 Met Val Arg Ser Asn Lys Gln Glu Pro Arg Val Ser Ser Thr Tyr Ile 1 5 10 15 Arg Ser Leu Val Lys Gln Gln Leu Ala Tyr Ser Thr Thr Met Thr Thr 20 25 30 Thr Thr Thr Thr Thr Thr Asn Asp Gly Ser Gly Gly Gly Lys Thr Gln 35 40 45 Thr Gln Thr His Lys Lys Gln Val Arg Arg Arg Leu His Thr Ser Arg 50 55 60 Pro Tyr Gln Glu Arg Leu Leu Asn Met Ala Glu Ala Arg Arg Glu Ile 65 70 75 80 Val Thr Ala Leu Lys Gln His Arg Ala Ser Met Arg Gln Ala Thr Arg 85 90 95 Ile Pro Pro Pro Gln Pro Pro Pro Pro Pro Gln Pro Leu Asn Leu Phe 100 105 110 Ser Pro Pro Pro Pro Pro Pro Pro Pro Asp Pro Phe Ser Trp Thr Asn 115 120 125 Pro Ser Leu Asn Phe Leu Leu Pro Asn Gln Pro Leu Gly Leu Asn Leu 130 135 140 Asn Phe Gln Asp Phe Asn Asp Phe Ile Gln Thr Ser Ser Thr Thr Ser 145 150 155 160 Ser Ser Ser Ser Ser Ser Thr Ser Ser Ser Ser Ser Ser Ile Phe Pro 165 170 175 Thr Asn Pro His Ile Tyr Ser Ser Pro Ser Pro Pro Pro Thr Phe Thr 180 185 190 Thr Ala Thr Ser Asp Ser Ala Pro Gln Leu Pro Ser Ser Ser Asn Gly 195 200 205 Glu Asn Asn Val Val Thr Ser Ala Trp Trp Ser Glu Leu Met Leu Lys 210 215 220 Thr Val Glu Pro Glu Ile Lys Pro Glu Thr Glu Glu Val Ile Val Val 225 230 235 240 Glu Asp Asp Val Phe Pro Lys Phe Ser Asp Val Met Glu Phe Pro Ser 245 250 255 Trp Leu Asn Gln Thr Glu Glu Glu Leu Phe His Pro Tyr Asn Leu Thr 260 265 270 Asp His Tyr Ser Ser Ser Pro His Asn Pro Pro Leu Ser Cys Met Glu 275 280 285 Ile Gly Glu Ile Glu Gly Met Asp Gly Asp Asp Trp Leu Ala 290 295 300 <210> 4 <211> 384 <212> PRT <213> Unknown <220> <223> TraesCS5B02G518900.1 amino acid <400> 4 Met Arg Asn Gln Arg Leu Ala Pro Ile Thr Cys Ser Ser Ser Lys Thr 1 5 10 15 Asn Asn Thr Lys Arg Glu Glu Pro His Leu Ser Gly Ala Tyr Ile Arg 20 25 30 Ser Leu Val Lys His Leu Ser Ser Ser Ser Thr Ala Arg Ser Lys Asp 35 40 45 His His His Ile Thr Met Gly Ser Lys Ser His Gln Glu Glu Gln Gln 50 55 60 Ala Pro Gln Thr Thr Pro Ser Ser Leu Gln Gln Gln Gln Pro His Lys 65 70 75 80 Lys Gln Val Arg Arg Arg Leu His Thr Ser Arg Pro Tyr Gln Glu Arg 85 90 95 Leu Leu Asn Met Ala Glu Ala Arg Arg Glu Ile Val Thr Ala Leu Lys 100 105 110 Ile His Arg Ala Ser Met Arg Glu Ala Lys Glu Gln Gln Gln His Gln 115 120 125 Gln Leu Val Gln Gln Leu Gln His Gln Gln Glu Val Gln Val Val Gln 130 135 140 Asp His Arg Val Ala Phe Ser Ala Pro Asn Met Ser Ser Tyr Gly Ser 145 150 155 160 Phe Ser Asp Tyr Leu His Asn Ser Tyr Pro Phe Glu His Ser Thr Ala 165 170 175 Pro Ser Asn Ser Gly Cys Ser Tyr Tyr Ser Ser Pro Pro Leu Leu Pro 180 185 190 Tyr His Thr Pro Val Val Ala Pro Met Val Pro Met Val Asp Ala Leu 195 200 205 Asp Gln Leu Leu Ser Leu Pro Thr Gln Pro Leu Gly Leu Asn Leu Thr 210 215 220 Phe Asp Gly Phe Asn Gly Gly Val Ala Ala Glu Asp Ala Lys Asn Cys 225 230 235 240 Thr Ala Thr Ser Pro Phe Asp Pro Pro Ser Leu Val Gln Gln Pro Ser 245 250 255 Pro Ala Ser Ser Tyr Ser Val Tyr Ser Ser Pro Pro Leu Ala Thr Met 260 265 270 Val Ser Gln Asp Met Ala Ser Ala Ala Ala Glu Asn Thr Ser Gln Leu 275 280 285 Leu His Arg Val Leu Asp Glu Glu Glu Met Ala Ala Ile His Ser Ala 290 295 300 Gly Glu Arg His Asp Ile Glu Trp Ser Asp Thr Val Asn Leu Ala Thr 305 310 315 320 Ser Ala Trp Trp Ser Arg Leu Leu Glu Ser Val Glu Gly Gly Glu Asp 325 330 335 Asp Asp Gly Ala Ala Ala Ala Gln Gln Thr Asn Thr Val Asp Ala Met 340 345 350 Gly Met His Leu Ser Asp Glu Cys Tyr Gly Gln Asp Val Ser Phe Pro 355 360 365 Cys Met Asp Ile Gly Glu Ile Glu Gly Trp Asp Ala Glu Trp Phe Ser 370 375 380 <210> 5 <211> 21 <212> DNA <213> Artificial Sequence <220> <223> 0067LP <400> 5 atccatgcct tcaatttctc c 21 <210> 6 <211> 21 <212> DNA <213> Artificial Sequence <220> <223> 0067RP <400> 6 ttcacacaag ccgtccttat c 21 <210> 7 <211> 19 <212> DNA <213> Artificial Sequence <220> <223> LB1.3 <400> 7 attttgccga tttcggaac 19 <210> 8 <211> 22 <212> DNA <213> Artificial Sequence <220> <223> AtSEP F <400> 8 cctccacttc atcttcctca tc 22 <210> 9 <211> 20 <212> DNA <213> Artificial Sequence <220> <223> AtSEP R <400> 9 cgcaggcttg ttgttatttt 20 <210> 10 <211> 30 <212> DNA <213> Artificial Sequence <220> <223> AtSEP_GFP F <400> 10 cctctagaat ggtcagatca aacaagcaag 30 <210> 11 <211> 29 <212> DNA <213> Artificial Sequence <220> <223> AtSEP_GFP R <400> 11 ccctcgagaa gcaagccagt cgtctccat 29 <210> 12 <211> 29 <212> DNA <213> Artificial Sequence <220> <223> TaSEP_GFP F <400> 12 aatctagaat gaggaatcaa agattagct 29 <210> 13 <211> 27 <212> DNA <213> Artificial Sequence <220> <223> TaSEP_GFP R <400> 13 aaggatccac gataaccact cctcatc 27 <210> 14 <211> 20 <212> DNA <213> Artificial Sequence <220> <223> beta-Tublin F <400> 14 aagatccgtg aagagtaccc 20 <210> 15 <211> 19 <212> DNA <213> Artificial Sequence <220> <223> beta-Tublin R <400> 15 gactgtgagg gaacggtac 19 <210> 16 <211> 32 <212> PRT <213> Unknown <220> <223> At5G21280 conserved sequences 1 <400> 16 Lys Gln Val Arg Arg Arg Leu His Thr Ser Arg Pro Tyr Gln Glu Arg 1 5 10 15 Leu Leu Asn Met Ala Glu Ala Arg Arg Glu Ile Val Thr Ala Leu Lys 20 25 30 <210> 17 <211> 25 <212> PRT <213> Unknown <220> <223> At5G21280 conserved sequences 2 <400> 17 Leu Asn Phe Leu Leu Pro Asn Gln Pro Leu Gly Leu Asn Leu Asn Phe 1 5 10 15 Gln Asp Phe Asn Asp Phe Ile Gln Thr 20 25 <110> REPUBLIC OF KOREA(MANAGEMENT: RURAL DEVELOPMENT ADMINISTRATION) <120> Plant growth regulatory genes encoding seeds plant specific extensin motif containing proteins and uses it <130> KPA200896-KR <160> 17 <170> KoPatentIn 3.0 <210> 1 <211> 2047 <212> DNA <213> Unknown <220> <223> At5G21280 nucleotide <400> 1 caaaagtaca cacagagaca ctgttagctc ttaagtaata tgtcatcgta agcaacaaaa 60 aaaaatgaaa gagcaactaa ttctcatttc tgataatatt ttctgatttt tcacaaaaat 120 aaaattctgt aaaattgggc tagagcagag tatttatatc acccacgagc ctttgcttct 180 tcttccttct tcatcacctc tcttcttctt tctgttctct ttccggagac atttgtctaa 240 atctcatcat cattcacaat ttactcaaac aaaaccttga gaaagagaaa aaaatgatcc 300 aagagaagaa gaaaaaccta ttaatgtctt ctcagtgaaa aaccataact aaaatcttta 360 atggtcagat caaacaagca agaaccacgc gtctcttcta cttacatccg atctctcgtg 420 aagcaacaac ttgcttattc caccactatg accaccacca ccaaacaac aacaaacgat 480 ggtagcggcg gcggaaaaac gcaaacgcaa acgcacaaga aacaagtgag gaggcgactt 540 cacacaagcc gtccttatca agaacgtctc ctaaacatgg ctgaagctcg tcgcgaaaatc 600 gtcaccgctt taaaacaaca tcgtgcttcc atgagacaag ccacaagaat cccaccgcct 660 caaccacccc caccaccgca gcctctaaat cttttctctc cgccgcctcc tcctcctcct 720 ccggatccat tttcttggac gaatccatct ctcaatttcc tcctccctaa tcaaccctta 780 gggttaaacc taaacttcca agatttcaac gacttcatcc aaacctcctc aacaacatct 840 tcatcatcat cttcctccac ttcatcttcc tcatcctcga tttttccgac gaatcctcat 900 atctactctt ccccttcacc tcctcccact ttcaccaccg ccacttcaga ttcagctcct 960 caactaccgt cttcctctaa tggagaaaac aacgtggtga cgtcagcttg gtggtcagaa 1020 ctcatgttga agacggtgga gccggagatt aaaccggaaa cggaagaagt catcgtagtc 1080 gaagatgacg tgtttccaaa gtttagtgac gtcatggaat ttccttcgtg gttaaaccaa 1140 acagaggaag aattatttca tccttataat ctcactgatc attactcatc atctcctcat 1200 aatcctccat tatcctggta attattttgt atttattttt cattacttag aattaaagtt 1260 attttgatcg gtgttgtttt tcctccggaa tcaccggtgg cgaaaattgc ggcggttctt 1320 gcataaactt cagtgattga gtttgagaca gaacacagaa aaaacagaat cgttcttttg 1380 tttggttttg ataattaaat cggagttatt ggattaaaaa tatgtgaaag cttgtggtat 1440 gagaaggatc cacataggtt tattgcacga tctgaagctg tccatttttt tcttttagta 1500 tatggtccaa tgataaataa tatcgccgac atctataaaa cactatcctt accccatttt 1560 tctgaactgt cccaaaaccg tcttgacgtc acccgatata aattatattt ttgttatgtt 1620 atcattgcac ttgcattatt caaatttggt tgttacttgt taggtaaatt atatatctct 1680 tttgtgtgtt ctaatgaaac atttttgatg tgtgatattc tttttacagt atggagatg 1740 gagaaattga aggcatggat ggagacgact ggcttgcttg aaccacaaag attgattact 1800 gttttttttt tcagacatga tgaatcaact ctttggttat tttcttatta tttttacctt 1860 attttattg tttttcttta taattttttc tttttgtttg gtgacatttt aatgaatctt 1920 taaaaataac aacaagcctg cggagagaaa atgttcgatt tggcaaagcc acatgaatga 1980 agtaaaactt tcattttcgt tgattgactt tgttttttgt taataactta ttattagact 2040 ttaaaatg 2047 <210> 2 <211> 1155 <212> DNA <213> Unknown <220> <223> TraesCS5B02G518900.1 nucleotide <400> 2 atgaggaacc agaggctagc tcctatcact tgctcctctt caaagaccaa caacaccaag 60 cgagaggagc cacacctctc cggggcttac attaggagcc ttgtgaagca tcttagctcc 120 tcatctacgg cgagatccaa agaccaccat cacatcacca tgggctccaa atcacaccaa 180 gaagagcaac aagccccaca aaccacacca tcatctctac agcaacagca gccacacaag 240 aagcaggtga ggaggaggct ccacacgagc aggccatatc aggagaggct actcaacatg 300 gcggaggcca ggcgagagat cgtcaccgct ctcaagatcc acagagcctc catgagagaa 360 gccaaggagc agcagcagca tcaacaactt gtgcaacaat tgcagcatca gcaagaagtc 420 caggtagtgc aagatcatag ggtagctttt agtgcaccca acatgagctc atatggttcc 480 ttctcagatt acttgcacaa tagttaccca ttcgaacact ccacagcacc aagcaacagt 540 ggttgctcat attattcatc tcctccactc ctcccttacc acacaccagt cgttgcaccc 600 atggttccca tggtagatgc tttagatcag ttgctgtcgc tgcctacgca gccactaggg 660 ctcaacctga cattcgatgg cttcaatggt ggtgttgctg ccgaagatgc caagaactgc 720 actgctacta gcccctttga tcctccttcg ttggtccaac aaccatcacc tgcctcctcc 780 tactctgtct actcctctcc accgctggcg acgatggtga gccaggacat ggcgtccgct 840 gctgccgaga acacctcaca gttgctgcac cgggtgctgg acgaggagga gatggcggcg 900 atccactcgg ccggggagcg gcatgacatc gagtggagcg acaccgtgaa cctggccacg 960 tcggcgtggt ggagcaggct actcgagagt gtggaaggcg gtgaagacga tgatggggcc 1020 gcggcggcgc agcagactaa cacggtggac gccatgggga tgcatctgag tgatgagtgc 1080 tacggccaag acgtgtcctt tccatgtatg gacatcggag agatcgaagg atgggacgcc 1140 gagtggttct catga 1155 <210> 3 <211> 302 <212> PRT <213> Unknown <220> <223> At5G21280 amino acid <400> 3 Met Val Arg Ser Asn Lys Gln Glu Pro Arg Val Ser Ser Thr Tyr Ile 1 5 10 15 Arg Ser Leu Val Lys Gln Gln Leu Ala Tyr Ser Thr Thr Met Thr Thr 20 25 30 Thr Thr Thr Thr Thr Asn Asp Gly Ser Gly Gly Gly Lys Thr Gln 35 40 45 Thr Gln Thr His Lys Lys Gln Val Arg Arg Arg Leu His Thr Ser Arg 50 55 60 Pro Tyr Gln Glu Arg Leu Leu Asn Met Ala Glu Ala Arg Arg Glu Ile 65 70 75 80 Val Thr Ala Leu Lys Gln His Arg Ala Ser Met Arg Gln Ala Thr Arg 85 90 95 Ile Pro Pro Pro Gln Pro Pro Pro Pro Pro Gln Pro Leu Asn Leu Phe 100 105 110 Ser Pro Pro Pro Pro Pro Pro Pro Pro Asp Pro Phe Ser Trp Thr Asn 115 120 125 Pro Ser Leu Asn Phe Leu Leu Pro Asn Gln Pro Leu Gly Leu Asn Leu 130 135 140 Asn Phe Gln Asp Phe Asn Asp Phe Ile Gln Thr Ser Ser Thr Thr Ser 145 150 155 160 Ser Ser Ser Ser Ser Ser Thr Ser Ser Ser Ser Ser Ser Ile Phe Pro 165 170 175 Thr Asn Pro His Ile Tyr Ser Ser Pro Ser Pro Pro Pro Thr Phe Thr 180 185 190 Thr Ala Thr Ser Asp Ser Ala Pro Gln Leu Pro Ser Ser Ser Asn Gly 195 200 205 Glu Asn Asn Val Val Thr Ser Ala Trp Trp Ser Glu Leu Met Leu Lys 210 215 220 Thr Val Glu Pro Glu Ile Lys Pro Glu Thr Glu Glu Val Ile Val Val 225 230 235 240 Glu Asp Asp Val Phe Pro Lys Phe Ser Asp Val Met Glu Phe Pro Ser 245 250 255 Trp Leu Asn Gln Thr Glu Glu Glu Leu Phe His Pro Tyr Asn Leu Thr 260 265 270 Asp His Tyr Ser Ser Ser Pro His Asn Pro Pro Leu Ser Cys Met Glu 275 280 285 Ile Gly Glu Ile Glu Gly Met Asp Gly Asp Asp Trp Leu Ala 290 295 300 <210> 4 <211> 384 <212> PRT <213> Unknown <220> <223> TraesCS5B02G518900.1 amino acid <400> 4 Met Arg Asn Gln Arg Leu Ala Pro Ile Thr Cys Ser Ser Ser Lys Thr 1 5 10 15 Asn Asn Thr Lys Arg Glu Glu Pro His Leu Ser Gly Ala Tyr Ile Arg 20 25 30 Ser Leu Val Lys His Leu Ser Ser Ser Ser Thr Ala Arg Ser Lys Asp 35 40 45 His His His Ile Thr Met Gly Ser Lys Ser His Gln Glu Glu Gln Gln 50 55 60 Ala Pro Gln Thr Thr Pro Ser Ser Leu Gln Gln Gln Gln Pro His Lys 65 70 75 80 Lys Gln Val Arg Arg Arg Leu His Thr Ser Arg Pro Tyr Gln Glu Arg 85 90 95 Leu Leu Asn Met Ala Glu Ala Arg Arg Glu Ile Val Thr Ala Leu Lys 100 105 110 Ile His Arg Ala Ser Met Arg Glu Ala Lys Glu Gln Gln Gln His Gln 115 120 125 Gln Leu Val Gln Gln Leu Gln His Gln Gln Glu Val Gln Val Val Gln 130 135 140 Asp His Arg Val Ala Phe Ser Ala Pro Asn Met Ser Ser Tyr Gly Ser 145 150 155 160 Phe Ser Asp Tyr Leu His Asn Ser Tyr Pro Phe Glu His Ser Thr Ala 165 170 175 Pro Ser Asn Ser Gly Cys Ser Tyr Tyr Ser Ser Pro Pro Leu Leu Pro 180 185 190 Tyr His Thr Pro Val Val Ala Pro Met Val Pro Met Val Asp Ala Leu 195 200 205 Asp Gln Leu Leu Ser Leu Pro Thr Gln Pro Leu Gly Leu Asn Leu Thr 210 215 220 Phe Asp Gly Phe Asn Gly Gly Val Ala Ala Glu Asp Ala Lys Asn Cys 225 230 235 240 Thr Ala Thr Ser Pro Phe Asp Pro Pro Ser Leu Val Gln Gln Pro Ser 245 250 255 Pro Ala Ser Ser Tyr Ser Val Tyr Ser Ser Pro Pro Leu Ala Thr Met 260 265 270 Val Ser Gln Asp Met Ala Ser Ala Ala Ala Glu Asn Thr Ser Gln Leu 275 280 285 Leu His Arg Val Leu Asp Glu Glu Glu Met Ala Ala Ile His Ser Ala 290 295 300 Gly Glu Arg His Asp Ile Glu Trp Ser Asp Thr Val Asn Leu Ala Thr 305 310 315 320 Ser Ala Trp Trp Ser Arg Leu Leu Glu Ser Val Glu Gly Gly Glu Asp 325 330 335 Asp Asp Gly Ala Ala Ala Ala Gln Gln Thr Asn Thr Val Asp Ala Met 340 345 350 Gly Met His Leu Ser Asp Glu Cys Tyr Gly Gln Asp Val Ser Phe Pro 355 360 365 Cys Met Asp Ile Gly Glu Ile Glu Gly Trp Asp Ala Glu Trp Phe Ser 370 375 380 <210> 5 <211> 21 <212> DNA <213> Artificial Sequence <220> <223> 0067LP <400> 5 atccatgcct tcaatttctc c 21 <210> 6 <211> 21 <212> DNA <213> Artificial Sequence <220> <223> 0067RP <400> 6 ttcacacaag ccgtccttat c 21 <210> 7 <211> 19 <212> DNA <213> Artificial Sequence <220> <223> LB1.3 <400> 7 attttgccga tttcggaac 19 <210> 8 <211> 22 <212> DNA <213> Artificial Sequence <220> <223> AtSEP F <400> 8 cctccacttc atcttcctca tc 22 <210> 9 <211> 20 <212> DNA <213> Artificial Sequence <220> <223> AtSEP R <400> 9 cgcaggcttg ttgttattt 20 <210> 10 <211> 30 <212> DNA <213> Artificial Sequence <220> <223> AtSEP_GFP F <400> 10 cctctagaat ggtcagatca aacaagcaag 30 <210> 11 <211> 29 <212> DNA <213> Artificial Sequence <220> <223> AtSEP_GFP R <400> 11 ccctcgagaa gcaagccagt cgtctccat 29 <210> 12 <211> 29 <212> DNA <213> Artificial Sequence <220> <223> TaSEP_GFP F <400> 12 aatctagaat gaggaatcaa agattagct 29 <210> 13 <211> 27 <212> DNA <213> Artificial Sequence <220> <223> TaSEP_GFP R <400> 13 aaggatccac gataaccact cctcatc 27 <210> 14 <211> 20 <212> DNA <213> Artificial Sequence <220> <223> beta-Tublin F <400> 14 aagatccgtg aagagtaccc 20 <210> 15 <211> 19 <212> DNA <213> Artificial Sequence <220> <223> beta-Tublin R <400> 15 gactgtgagg gaacggtac 19 <210> 16 <211> 32 <212> PRT <213> Unknown <220> <223> At5G21280 conserved sequences 1 <400> 16 Lys Gln Val Arg Arg Arg Leu His Thr Ser Arg Pro Tyr Gln Glu Arg 1 5 10 15 Leu Leu Asn Met Ala Glu Ala Arg Arg Glu Ile Val Thr Ala Leu Lys 20 25 30 <210> 17 <211> 25 <212> PRT <213> Unknown <220> <223> At5G21280 conserved sequences 2 <400> 17 Leu Asn Phe Leu Leu Pro Asn Gln Pro Leu Gly Leu Asn Leu Asn Phe 1 5 10 15 Gln Asp Phe Asn Asp Phe Ile Gln Thr 20 25
Claims (19)
상기 식물체 생장 조절은 다음 중 선택되는 어느 하나 이상의 특징을 가지는 것인, 조성물:
i) 식물체의 지방산 함량 조절;
ii) 식물체 내 글리옥시좀 또는 퍼옥시좀 발현 조절;
iii) 식물세포의 크기 조절;
iv) 식물체 내 지질 분해 조절;
v) 식물체 내 글리옥실산 회로 관련 유전자인 TAG 모빌라이제이션 (SDP1), 베타 산화 (PXA1, ACX2-4, KAT2), 글리옥실산 회로 (CSY3, ACO3, MLS, PMDH1) 또는 퍼옥시좀 생성 (PEX11A, D, E) 유전자 발현 조절;
vi) 식물체의 개화시기 조절; 및
vii) 식물체의 산화스트레스 저항성 조절.
A composition for controlling plant growth comprising a sep gene encoding the amino acid sequence of SEQ ID NO: 4,
A composition in which the plant growth control has one or more characteristics selected from the following:
i) Control of fatty acid content in plants;
ii) regulation of glyoxisome or peroxisome expression in plants;
iii) regulation of plant cell size;
iv) regulation of lipid degradation in plants;
v) Genes related to the glyoxylate cycle in plants: TAG mobilization (SDP1), beta oxidation (PXA1, ACX2-4, KAT2), glyoxylate cycle (CSY3, ACO3, MLS, PMDH1) or peroxisome production ( PEX11A, D, E) regulation of gene expression;
vi) Control of flowering time of plants; and
vii) Control of oxidative stress resistance in plants.
The composition according to claim 1, wherein the sep gene is derived from wheat.
The composition according to claim 2, wherein the sep gene is represented by the base sequence of SEQ ID NO: 2.
The composition of claim 1, wherein the composition comprises a recombinant vector for plant growth control containing a sep gene encoding the amino acid sequence of SEQ ID NO: 4.
상기 식물체 생장 조절은 다음 중 선택되는 어느 하나 이상의 특징을 가지는 것인, 벡터:
i) 식물체의 지방산 함량 조절;
ii) 식물체 내 글리옥시좀 또는 퍼옥시좀 발현 조절;
iii) 식물세포의 크기 조절;
iv) 식물체 내 지질 분해 조절;
v) 식물체 내 글리옥실산 회로 관련 유전자인 TAG 모빌라이제이션 (SDP1), 베타 산화 (PXA1, ACX2-4, KAT2), 글리옥실산 회로 (CSY3, ACO3, MLS, PMDH1) 또는 퍼옥시좀 생성 (PEX11A, D, E) 유전자 발현 조절;
vi) 식물체의 개화시기 조절; 및
vii) 식물체의 산화스트레스 저항성 조절.
A recombinant vector for plant growth control containing a sep gene encoding the amino acid sequence of SEQ ID NO: 4,
The plant growth control vector has one or more characteristics selected from the following:
i) Control of fatty acid content in plants;
ii) regulation of glyoxisome or peroxisome expression in plants;
iii) regulation of plant cell size;
iv) regulation of lipid degradation in plants;
v) Genes related to the glyoxylate cycle in plants: TAG mobilization (SDP1), beta oxidation (PXA1, ACX2-4, KAT2), glyoxylate cycle (CSY3, ACO3, MLS, PMDH1) or peroxisome production ( PEX11A, D, E) regulation of gene expression;
vi) Control of flowering time of plants; and
vii) Control of oxidative stress resistance in plants.
상기 식물체 생장 촉진은 다음 중 선택되는 어느 하나 이상의 특징을 가지는 것인, 식물체의 생장 촉진 방법:
i) 식물체의 지방산 함량 감소;
ii) 식물체 내 글리옥시좀 또는 퍼옥시좀 발현 증가;
iii) 식물세포의 크기 증가;
iv) 식물체 내 지질 분해 증가;
v) 식물체 내 글리옥실산 회로 관련 유전자인 TAG 모빌라이제이션 (SDP1), 베타 산화 (PXA1, ACX2-4, KAT2), 글리옥실산 회로 (CSY3, ACO3, MLS, PMDH1) 또는 퍼옥시좀 생성 (PEX11A, D, E) 유전자 발현 증가;
vi) 식물체의 개화시기 촉진; 및
vii) 식물체의 산화스트레스 저항성 증가.
A method for promoting the growth of a plant, comprising the step of overexpressing the sep gene encoding the amino acid sequence of SEQ ID NO: 4 in the plant,
A method of promoting plant growth, wherein the plant growth promotion has one or more characteristics selected from the following:
i) reducing the fatty acid content of the plant;
ii) increased expression of glyoxysomes or peroxisomes in plants;
iii) increase in size of plant cells;
iv) increased lipid breakdown in the plant;
v) Genes related to the glyoxylate cycle in plants: TAG mobilization (SDP1), beta oxidation (PXA1, ACX2-4, KAT2), glyoxylate cycle (CSY3, ACO3, MLS, PMDH1) or peroxisome production ( PEX11A, D, E) Increased gene expression;
vi) Promoting the flowering time of plants; and
vii) Increased resistance to oxidative stress in plants.
The method of claim 7, wherein the plants include Arabidopsis thaliana, tobacco, eggplant, pepper, tomato, burdock, mugwort, lettuce, bellflower root, spinach, Swiss chard, sweet potato, celery, carrot, water parsley, parsley, Chinese cabbage, cabbage, pollack radish, watermelon, Promoting plant growth, which is at least one selected from the group consisting of melon, cucumber pumpkin, gourd, strawberry, soybean, mung bean, kidney bean, pea, rice, barley, wheat, rye, corn, sugarcane, oats, and onion. method.
The method of claim 7, wherein the sep gene is represented by the base sequence of SEQ ID NO: 2.
The method of claim 7, wherein the promoting plant growth restores hypocotyl length, leaf area, flowering time, or epidermal cell length to the level of wild species.
상기 식물체 생장 저해는 다음 중 선택되는 어느 하나 이상의 특징을 가지는 것인, 식물체의 생장 저해 방법:
i) 식물체의 지방산 함량 증가;
ii) 식물체 내 글리옥시좀 또는 퍼옥시좀 발현 감소;
iii) 식물세포의 크기 감소;
iv) 식물체 내 지질 분해 감소;
v) 식물체 내 글리옥실산 회로 관련 유전자인 TAG 모빌라이제이션 (SDP1), 베타 산화 (PXA1, ACX2-4, KAT2), 글리옥실산 회로 (CSY3, ACO3, MLS, PMDH1) 또는 퍼옥시좀 생성 (PEX11A, D, E) 유전자 발현 조절;
vi) 식물체의 개화시기 지연; 및
vii) 식물체의 산화스트레스 저항성 감소.
A method of inhibiting the growth of a plant, comprising the step of inhibiting the expression of the sep gene encoding the amino acid sequence of SEQ ID NO: 4 in the plant,
A method of inhibiting plant growth, wherein the plant growth inhibition has one or more characteristics selected from the following:
i) increasing the fatty acid content of the plant;
ii) Reduced expression of glyoxysomes or peroxisomes in plants;
iii) Reduction in size of plant cells;
iv) reduction of lipid breakdown in plants;
v) Genes related to the glyoxylate cycle in plants: TAG mobilization (SDP1), beta oxidation (PXA1, ACX2-4, KAT2), glyoxylate cycle (CSY3, ACO3, MLS, PMDH1) or peroxisome production ( PEX11A, D, E) regulation of gene expression;
vi) delay in flowering of plants; and
vii) Reduction of oxidative stress resistance in plants.
The method of claim 12, wherein the plants include Arabidopsis thaliana, tobacco, eggplant, pepper, tomato, burdock, mugwort, lettuce, bellflower root, spinach, Swiss chard, sweet potato, celery, carrot, water parsley, parsley, Chinese cabbage, cabbage, pollack radish, watermelon, Inhibition of plant growth, which is one or more selected from the group consisting of melon, cucumber pumpkin, gourd, strawberry, soybean, mung bean, kidney bean, pea, rice, barley, wheat, rye, corn, sugarcane, oats, and onion. method.
The method of claim 12, wherein the sep gene is represented by the base sequence of SEQ ID NO: 2.
The method of claim 12, wherein the plant growth inhibition reduces hypocotyl length, leaf area, flowering time, or epidermal cell length compared to the level of wild species.
The composition for regulating plant growth according to claim 1, wherein the plant growth control involves controlling hypocotyl length, leaf area, flowering time, or epidermal cell length.
The method of claim 1, wherein the plants include Arabidopsis thaliana, tobacco, eggplant, pepper, tomato, burdock, mugwort, lettuce, bellflower root, spinach, Swiss chard, sweet potato, celery, carrot, water parsley, parsley, Chinese cabbage, cabbage, pollack radish, watermelon, A plant that is one or more selected from the group consisting of melon, cucumber pumpkin, gourd, strawberry, soybean, mung bean, kidney bean, pea, rice, barley, wheat, rye, corn, sugar cane, oats, and onion.
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GenBank Accession Number KAF7069115 (2020.08.20.) 1부.* |
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